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^"L 



PROCEEDINGS 



OF THE 



aENERAL MEETmaS FOR SCIENTIFIC BUSINESS 



ZOOLOGICAL SOCLETY 

OF LONDON. 
1911, pp. 1-555, 

WITH 16 Plates and 144 Text-figures, 



PRINTED FOR THE SOCIETY, 
AND SOLD AT THEIE HOUSE IN REGENT'S PARK". 

LONDON : 

MESSRS. LONGMANS, GREEN, AND CO., 

PATERNOSTER RO'iV. 



1\ 



zzs\> 



LIST 



OF THE 



COUNCIL AND OFFICERS 

OF THE 

ZOOLOGICAL SOCIETY OF LONDON. 

1911. 



COUNCIL. 
His Grace The Duke of Bedford, K.G., President. 



The Earl of Altamont, F.S.A. 
Prof. J. Rose Bradford, M.D., 

D.Sc, F.R.S., Vice-President. 
Lt.-Col. Sir R. Havelock- 

Charles, K.C.V.O., M.D. 
Alfred H. Cocks, Esq., M.A, 
The Rt. Hon. the Earl of 

Cromer, P.C, G.C.B., 

G.C.M.G. 
F. D. Dawtrey Drbwitt, Esq., 

M.A., M.D. 
Charles Drummond, Esq., 

Treasurer. 
Sir Edward Dtjrand, Bt., C.B. 
Frederick Gillett, Esq., Vice- 
President. 
Sidney F. Harmer, Esq., M.A., 

Sc.D., F.R.S., Vice-President. 



Sir Walter Roper Lawrence, 

Bt., G.C.I.E. 
Sir Edmund G. Loder, Bt. 
E. G. B. Meade-Waldo, Esq., 

Vice-President. 
P. Chalmers Mitchell, Esq,, 

M,A., D.Sc, Hon. LL.D., 

F.R.S. , Secretary. 
W. R. Ogilvie-Grant, Esq. 
Adrian D. W. Pollock, Esq. 
A. Trevor - Battye, Esq., 

M.A. 
Anthony H. Wingfield, 

Esq. 
A. Smith Woodward, Esq., 

Ijh.Jy. ,Y.'K.^.,Vice-President. 
Henry WooDWARD,EsQ,, LL.D., 

F.R.S., Vice-President. 



PRINCIPAL OFFICERS. 

P. Chalmers Mitchell, M.A., D.Sc, Hon.LL.D., F.R.S., 

Secretary. 
Frank E. Beddaed, M.A., F.R.S., Prosector. 
R. I. Pocock, F.R.S., F.L.S., Curator of Mammals, and 

Resident Superintendent of the Gardens. 
D. Seth-Smith, Curator of Bird^s and Inspector of Works, 
Henry G. Plimmer, F.R.S., M.R.C.S,, Pathologist. 
F. H. Waterhouse, Librarian. 
John Barrow, Accountant, 
W. H. Cole, Chief Clerk. 



LIST OF CONTEMS. 

1911, pp. 1-555 



EXHIBITIONS AND NOTICES. 

Page 
The Secretary. Eeport on the Additions to the Society's 
Menagerie during the months of June, July, August, 
September, and October, 1910 1 

Mr. R. I. PococK,. F.L.S., F.Z.S. Exhibition of a living 

specimen of a Black Rat (J/«g r«<i?6s) from Sark 4 

Mr. R. I. PocoGK, F.L.S., F.Z.S. Exhibition of a female 
hybrid between a male Black Lemur [Lemur macaco) 
and a female of the Red-fronted variety of the Fulvous 
Lemur [Lemur fidvus rujifrons) 5 

The Hon, N. Charles Rothschild, M. A., F.Z.S. Exhibition 
of some Fleas interesting in connection with the 
spreading of plague 5 

Mr. J. Lewis Bonhotb, M.A., F.L.S., F.Z.S. Exhibition 
of, and remarks upon, a young Cairo Spiny Mouse 
[Acomys cahirinns) about twelve hours old , 5 

Mr. J. Lewis Bonhote, M.A., F.L.S., F.Z.S. Exhibition of , 
and remarks upon, a pair of hybrids between the 
Bramble-Finch [Fringilla montifringilla) and the 
Chaffinch [Fi^ingilla ccelebs) 6 

Mrs. R. Haig Thomas, F.Z.S. Exhibition of a series of skins 
illustrating an experiment in Pheasant-breeding, and 
an account of the production of Thaumalea obsoura in 
the F„ generation from a ci-oss between T. amhersti ^ 
and t. picta <S 6 



IV 

Page 
Dr. W. KicoLL and Prof. E. A. Minchin, M.A., V.P.Z.S. 
Exhibition of, and remarks upon, two species of 
Cysticei'coids from the Jint-Q.eH.^Ceratoph'i/llusfasciatua). 
(Text-figs. 1 & 2) 9 

Dr. William Nicoll. Demonstration of his method for the 

collection of Trematodes 45 

Dr. R. T. Leiper, F.Z.S. Exhibition of photographs and 
specimens showing the Nematode infection known as 
Onchocei-ciasis 45 

Dr. J. F. Gemmill, M.A.. D.Sc. Notice of a paper on "The 

Development of Solaster endeca Forbes " 45 

Mr. D. Seth-Smith, F.Z.S. Exhibition of, and remarks 
\ipon, living examples of the Australian Budgerigar 
or Undulated Grass- Parrakeet(i/efoj9si^^«ct6S undulatus), 
showing three colour-phases 45 

The Secretary. Report on the Additions to the Society's 

Menagerie during the month of November 1910 100 

Dr. H. Hammond Smith, M.R.C.S., F.Z.S. Exhibition of, 
and remarks upon, a specimen of the Red Grouse 
{Lagopus scoticus) which displayed a curious variety 
of the ordinary plumage 1 00 

Mr. D. Seth-Smith, F.Z.S. Exhibition of some skins of the 
Austi'alian Yellow-rumped Finch {^Munia flaviprymrua) 
showing markings tending towards those of M. castanei- 
thorax 101 

Dr. W. E. Hoyle, M.A., F.Z.S. Explanation of the Report 
of the International Commission on Zoological Nomen- 
clature presented to the Gi'az Meeting of the 
International Zoological Congress 101 

The Secretary. Report on the Additions to the Society's 

Menagerie during the month of December 1910 181 

Dr. C. W. Andrews, F.R.S., F.Z.S. Exhibition of the 
skull of a Sabre-toothed Tiger (Smilodoti calif or nicies), 
from an asphalt deposit in California 181 

The Secretary. Exhibition of a mounted specimen of the 

Plat}qD\is ( Ornithorhynchns paradoxus) 181 



V 

Page 
Mr. Edward Gerrard, F.Z.S. Exhibition of the head of a 
Caribou {^Rangifer tarandus caribou) with abnonnal 
antlers. Also the heads of an Eland {Tait7'otrayus 
derhianus gigas), and of three White-tailed Deer {Dorc- 
elaphus americanus macri(,rtis). (Text-fig. 52) 181 

The Secretary. Report on the Additions to the Society's 

Menagerie during the month of January 1911 314 

Dr. H. Hammond Smith, M.R.O.S., F.Z.S. Exhibition of 

three skins of male Pheasants assuming female plumage 314 

Mr. G. A. BouLENGER, F.R.S., V.P.Z.S. Notice of a paper 
"On a Collection of Fishes from the Lake Ngami 
Basin, Bechuanaland " 314 

Mr. Alfred H. Cocks, M.A., F.Z.S. Exhibition of photo- 
graphs of, and remarks upon, the female Brindled Gnu 
recently born in the Society's Gardens. (Text-figs. 97 
& 98) 314 

The Secretary. Exhibition of a series of lantern-slides 
illustrating some of the most remarkable features of 
Mr. Carl Hagenbeck's new Tierpark at Stellingen 358 

Mr. Ernest C. Oberholtzer. Exhibition of lantern- slides 
and photographs illustrating an account of some 
Obsei'vations on Moose 358 



PAPERS. 



Page 

1 . On the Inheritance of the Webf oot Character in Pigeons. 

By J. Lewis Bonhote, M.A., F.L.S., F.Z.S. (Text- 
tigs. 3 & 4.) 14 

2. Kotes on the little-known Lizard Lacerta jacJcsoni Blgr., 

W'ifch Special Reference to its Cranial Characters. By 
Edward Degen, F.Z.S. (Text-figs. 5-7.) 19 

3. On the Peloponnesian Lizard (Lacerta peloponnesiaca 

Bibr.). By G. A. Boulengee, F.R.S., V.P.Z.S. 
(Plate L and Text-fig. 8.) 37 

4. Remarks on Two Species of Fishes of the Genus Gobius, 

from Observations made at RoscofF. By Edwaed G. 

BOULENGER 40 

5. On a Possible Cause of Pneumo-enteritis in the Red 

Grouse {Lagopus scoticus). By H. B. Fantham, D.Sc, 
B.A.. F.Z.S., and H. Hammond Smith, M.R.C.S., 
L.R.C.P., F.Z.S 46 

6. On the Alimentary Tract of certain Birds and on the 

Mesenteric Relations of the Intestinal Loops. By 
Frank E, Beddard, M.A., F.R.S., F.Z.S., Prosector to 
the Society. (Text-figs. 9-28.) 47 

7. On the Specimens of Spotted Hysenas in the British 

Museum (Natural History). By Prof. Angel Cabrera, 
C.M.Z.S 93 

8. On the Segmentation of the Occipital Region of the Head 

in the Batrachia Urodela. By Edwin S. Goodrich, 
M.A., F.R.S., F.Z.S., Fellow of Merton College, Oxford. 
(Text-figs. 29-5L) 101 

9. The Mammals of the Tenth Edition of Linnaeus ; an 

Attempt to fix the Types of the Genera and the exact 
Bases and Localities of the Species. By Oldfield 
Thomas, F.R.S., F.Z.S 120 



10. The Duke of Bedford's Zoological Exploration of Eastern 

Asia.— XIII. On Mammals from the Provinces of 
Kan-su and Sze-chwan, Western China. By Oldfield 
Thomas, F.R.S., F.Z.S 158 

11. On the Structure and Function of the Gas Glands and 

Retia Mirabilia associated with the Gas Bladder of 
some Teleostean Fishes, with Notes on the Teleost 
Pancreas. By W. N. F. Woodlakd, F.Z.S. , The 
Zoological Department, University College, London. 
(Plates II.-IX. and Text-figs. 53-62.) 183 

12. On Skulls of Oxen from the Roman Military Station at 

Newstead, Melrose. By J. C. Ewart, M.D., F.R.S., 
F.Z.S. (Text-figs. 63-91.) 249 

13. Plankton from Christmas Island, Indian Ocean. — I. On 

Copepoda of the Family Coryceeida?. By George P. 
Farran. (Plates X.-XIV.) 282 

14. On some New Zealand Spiders. Bv H. R. Hogg, M.A., 

F.Z.S. (Text-figs. 92-96.) " 297 

15. Report on the Deaths which occurred in the Zoological 

Gardens during 1910. By H. G. Plimmer, F.R.S., 
"T.Z.S., Pres. R.M.S., Pathologist to the Society 318 

16. A Contribution to the Study of the Variations of the 

Spotted Sala,mander (Salamandra maculosa). By 
Edward G. Boulenger. (Plate XV. and Text- 
figs. 99-102.) 323 

17. On the Mountain Nyala, Tragelaphus buxtoni. By R. 

Lydekker. (Plate XVI. and Text-fig. 103.) 348 

18. Observations on diflerent Gibbons of the Genus ZTv/ZoSft^es 

now or recently living in the Society's Gardens, and on 
Symphalangihs syndactylus, with Notes on Skins in the 
Natural History Museum, S. Kensington.. By Dr. F. 

D. Welch, F.Z.S 353 

19. Some New Siphonaptera from China. By Karl Jordan, 

Ph.D., F.E.S., and the Hon, N.. Charles Roth- 
schild, M.A., F.Z.S., F.E,S. (Text-figs. 104-124.) ... 365 

20. Contributions to the Anatomy of the Anura. By Frank 

E. Beddard, M.A., F.R.S., F.Z.S., Prosector to the 
Society. (Text-figs.- 125-133.) 393 



Vlll 

Page 

21. On the Spei'iiiatophores in Earthworms of the Genus 

Pheretima ( = Perichceta). By Frank E. Beddard, M. A. , 
F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 
] 34-136.)... ,.. 412 

22. A Rare Beaked Whale. By R. Lydekkek. (Text-figs. 

137-139.) 420 

23. Age- Phases of the RorquaL By R. Lydekker 423 

24. On Longevity and Relative Viability in Mammals and 

Birds .; with a Note on the Theorj'" of Longevity. By 
P. Chalmers Mitchell, M.A., D.Sc, LL.D., F.R.S., 
Secretary to the Society 425 

25. Some new Parasitic Nematodes from Tropical Afi'ica, 

By Robert T. Leiper, D.Sc, M.B., F.Z.S. (Text-figs. 
140-144.) 549 



ADDENDUM. 

A correction to Dr, R. T, Leiper's Memoir on the " Entozoa 

of the Hippopotamus " - . 555 



ALPHABETICAL LIST 



CON TRIBUTORS, 

With References to the several Articles contributed by each. 
(1911, pp. 1-555.) 



Page 
Andrews, Charles William, B.A., D.Sc, F.R.S., F.Z.S. 

Exhibition of the sknll of a Sabre-toothed Tiger 
(Smilodon califomicus), from an asphalt deposit in Cali- 
fornia 181 



Beddard, Frank E., M.A., F.R.S., F.Z.S., Prosector to the 
Society. 

On the Alimentary Tract of certain Birds and on the 
Mesenteric Relations of the Intestinal Loops. (Text- 
figs. 9-28) 47 

Contributions to the Anatomy of the Anura. (Text- 
figs. 125-133.) 393 

On the Spermatophores in Earthworms of the Genus 
Pheretima {=Perichceta). (Text-figs. 134-136.) 412 



X 

Page 
BoNHOTE, J. Lewis, M.A., F.L.S., F.Z.S. 

Exhibition of", and remarks upon, a. young Cairo Spiny 
Mouse (^Acomys cahirimos) about twelve hours old 5 

Exhibition of, and remarks upon, a pair of hybrids 
between the Bramble-Finch (Fringilla montifringilla) and 
the Chaffinch {Fringilla coelehs) 6 

On the Inheritance of the Webfoot Character in 
Pigeons. (Text-figs. 3 & 4.) 14 

BOCJLENGER, EdWARD G. 

Remarks on Two Species of Fishes of the Genus 
Gohius, from Observations made at Roscoff 40 

A Contribution to the Study of the Variations of the 
Spotted Salamander (Salcimandra uiacidosa). (Plate XY. 
and Text-figs. 99-102.) 323 



BouLENGER, George A., F.R.S., y.P.Z.S. 

On the Peloponnesian Lizard {Lacerta p6lo2)onnesiaca 
Bibr.). (Plate L and Text-fig. 8.) 37 

Notice of a paper " On a Collection of Fishes from the 
Lake Ngami Basin, Bechuanaland " 314 



Cabrera, Prof. Angel, C.M.Z.S. 

On the Specimens of Spotted Hyaenas in the British 
Museum (Natural History) 9'i 



Cocks, Alfred H., M.A., F.Z.S. 

Exhibition of photographs of, and remarks upon, the 
female Brindled Gnu recently born in the Society's 
Gardens. (Text-figs. 97 & 98.) 314 



XI 

Pago 
Degen, Edward, F.Z.S. 

Notes on the little-known Lizai-d Laceriajacksoni B\gr., 
with Special Reference to its Cranial Chai-acters. (Text- 
figs. 5-7.) 19 

EwART, James Cossar, M.D., F.R.S., F.Z.S. 

On Skulls of Oxen from the Roman Military Station 
at ISTewsteacl, Melrose. (Text-figs. 63-91.) 249 

Fantham, Harold B., D.Sc, B.A., F.Z.S. , and Smith, 
H. Hammond, M.R.C.S., L.R.O.P., F.Z.S. 

On a Possible Cause of Pneumo-enteritis in the Red 
Grouse [Lagojnis scoticus) 46 

Farran, George P. 

Plankton from Christmas Island, Indian Ocean. — I. 
On Copepoda of the Family Corycseidfe. (Plates X.-XIY.) 282 

Gemmill, Dr. James F., M.A., D.Sc, F.Z.S., Lecturer on 
Embryology in the University of Glasgow. 

Notice of a paper on " The Development of Solaste7' 
endeca Forbes " 45 

Gerrard, Edward, F.Z.S. 

Exhibition of the head of a Caribou {Rangifer tarandus 
caribou) with abnormal antlers. Also the heads of an 
Eland {Tatirotragus derhiamis gigas), and of three White- 
tailed Deer (Dorcelaphus mnericamts macrurus). (Text- 
fig. 52.) 181 

Goodrich, Edwin S., M.A., F.R.S., F.Z.S., Fellow of 
Merton College, Oxford. 

On the Segmentation of the Occipital Region of the 
Head in the Batrachia Urodela, (Text-figs. 29-51.) ... 101 



Xll 

Hogg, Henry R,, M.A., F.Z.S. 

On some New Zealand Spiders. (Text-figs. 92-96.) ... 297 

HoYLE, Dr. William E., M.A., F.Z.S., English Member of 
the International Commission on Zoological Nomen- 
clature. 

Explanation of the Report of the International Com- 
mission on Zoological Nomenclatui-e presented to the 
Graz Meeting of the International Zoological Congress... 101 

JoEDAN, Karl, Ph.D., F.E.S., and Rothschild, The Hon. 
N. Charles, M.A., F.Z.S., F.E.S. 

Some New Siphonaptera from China. (Text-figs. 
104-124.) 365 

Leiper, Robert T., D.Sc, M.B., F.Z.S. 

Exhibition of photographs and specimens showing the 
Nematode infection known as Onchocerciasis 45 

Some new Parasitic Nematodes from Tropical Africa. 
(Text-figs. 140-144.) 549 

A correction to the Memoir on the " Entozoa of the 
Hippopotamus " 555 

Lydekker, Richard. 

On the Mountain Nyala, Tragelaphtts huxtoni. (Plate 
XYI. and Text-fig. 103.) 348 

A Rare Beaked Whale. (Text-figs. 137-139.) 420 

Age-Phases of the Rorqual 423 

MmcHiN, Prof. Edward A., M.A., V.P.Z.S., and Nicoll, 
Dr. William, F.Z.S. 

Exhibition of, and remarks upon, two species of Cysti- 
cercoids from the Rat-flea {Geratophylhis fasciatus). 
(Text-figs. 1 & 2.) 9 



XIU 

Page 
Mitchell, P. Chalmers, M.A., D.Sc, Hon. LL.D., F.R.S., 

F.Z.S., Secretary to the Society. 

Report on the Additions to the Society's Menagerie 
during the months of June, Jvily, August, September, 
and October, 1910 1 

Report on the Additions to the Society's Menagerie 
during the month of November 1910 100 

Report on the Additions to the Society's Menagerie 
during the month of December 1910 181 

Exhibition of a mounted specimen of the Platypus 
{Ornithorhynchus paradoxus) 181 

Report on the Additions to the Society's Menagerie 
during the month of January 1911 314 

Exhibition of a series of lantei-n-slides illustrating 
some of the most remarkable features of Mr. Carl 
Hagenbeck's new Tierpark at Stellingen 358 

On Longevity and Relative Viability in Mammals and 
Birds; with a ISTote on the Theory of Longevity 425 



NicoLL, Dr. William, F.Z.S., Lister Institute of Preventive 
Medicine. 

Demonstration of his method for the collection of 
Trematodes 45 



NicoLL, Dr. William, F.Z.S., and Minchin, Prof. E. A., 
M.A., V.P.Z.S. 

Exhibition of, and remarks upon, two species of Cysti- 
cercoids from the Rat-flea [Ceratophyllus fasciatus). 
(Text-figs. 1& 2.) 



XIV 

Page 
Oberholtzer, Ernest C. 

Exhibition of lantern-slides and photograph, illus- 
trating an account of some Observations on Moose 358 



Pltmmer, Henry G., F.R.S., F.Z.S., Pres.R.M.S., Patho- 
logist to the Society. 

Report on the Deaths which occurred in the Zoological 
Gardens durinsf 1910 : 318 



PococK, Reginald I., F.R.S., F.L.S., F.Z.S., Curator of 
Mammals, and Resident Superintendent of the 
Gardens. 

Exhibition of a living specimen of a Black Rat [Mus 
rattvs) from Sark 4 

Exhibition of a female hybrid between a male Black 
Lemur {Lemur macaco) and a female of the Red-fronted 
variety of the Fulvous Lemur [Lemur fuhms rujjifrons)... 5 

Rothschild, The Hon. N. Charles, M.A., F.Z.S., F.E.S. 

Exhibition of some Fleas interesting in connection 
with the spreading of plague 5 

Rothschild, The Hon. N. Charles, M.A., F.Z.S., F.E.S., 
and Jordan, Karl, Ph.D., F.E.S. 

Some New Siphonaptera from China. (Text-figs. 104- 
124.) 365 

Sbth-Smith, D., F.Z.S., Curator of Birds. 

Exhibition of, and remarks upon, living examples of 
the Australian Budgerigar or Undulated Grass- Parrakeet 
' {Melopsittacus imdulatus), showing thi-ee colour-phases ... 45 



Pngo 
Seth-Smith, D., F.Z.S. {Gontinued.) 

Exhibition of some skins of tiie Australian Yellow- 
rumped Finch {Iftmia flaviprymna) showing markings 
tending towards those of M. castaneithorax 101 



Smith, H. Hammond, M.R.C.S., L.R.C.P., F.Z.S. 

Exhibition of, and remarks upon, a specimen of the 
Red Grouse {Lagopus scoticus) which displayed a curious 
variety of the ordinary plumage 100 

Exhibition of three skins of male Pheasants assuming 
female plumage 314 



Smith, H. Hammond, M.R.O.S., L.R.C.P., F.Z.S., and 
Fantham, Harold B., D.Sc, B.A., F.Z.S, 

On a Possible Cause of Pneumo-enteritis in the Red 
Grouse (Lagopios scoticus) 46 



Thomas, Oldpield, F.R.S., F.Z.S. 

The Mammals of the Tenth Edition of Linnseus ; an 
Attempt to fix the Types of the Genera and the exact 
Bases and Localities of the Species 120 

The Duke of Bedford's Zoological Exploration of 
Eastern Asia. — XIII. On Mammals from the Provinces 
of Kan-su and Sze-chwan, Western China 158 



Thomas, Mrs. R. Haig, F.Z.S. 

Exhibition of a series of skins illustrating an experi- 
ment in Pheasant-breeding, and an account of the 
production of Tliawnalea ohscura in the F^ generation 
from a cross between T, amhersti $ and T, picta <S ^ 



Page 
Welch, Dr. Frederick D., F.Z.S. 

Observations on different Gibbons of the Genus Hylo- 
bates now or recently living in the Society's Gardens, and 
on Symphalangus syndactylus, with Notes on Skins in 
the Natural History Museum, S. Kensington 353 



Woodland, William N. F,, F.Z.S., The Zoologicri Depart- 
ment, University College, London. 

On the Structure and Function of the Gas Glands and 
Retia Mirabilia associated with the Gas Bladder of some 
Teleostean Fishes, with Notes on the Teleost Pancreas. 
(Plates II. -IX. and Text-figs. 53-62.) 183 



LIST OF PLATES. 

1911, pp. 1-555. 



Plate Pao^e 

I. Lacerta peloponnesiaoa , 37 

III. I 

IV. I 

V. ;^.Gas Glands of Teleoslei 183 

VI. I 

VII. j 

VIII. I 

IXJ 

X. 1-6. Corycella brevis. 7-11. C. curtn -^ 

XI. 1-6. Corycella curia. 7. C. brevis. 8, 9. Corycella sp. J . I 

10. C. carinata. 11, 12. Coryc<si(,s gracilicaudatus. I 

XII. 1-8. Corycfsus catus. 4-6. C. obtusus. 7. C. dubius. ' 

8, 9. C. tenuis ^^"^ 

XIII. 1-6. Corycceus murrayi. 7-9. C. andrewsi 

XIV. 1-4. Corycceus andrewsi. 5-9. C. dubius j 

XV. Salamandra maculosa var. tceniata, var. molleri, and forma 

typica 323 

XVI. The Mountain Nyala, Tracjdnphvs buxfoni 348 



Proc. Zool. Soc, — 1911, 



LIST OF TEXT-FIG UTIE8. 

1911, pp. 1-555. 



Page 

1. Cysticercoid of Hymenolepis diminuta 10 

2. Cysticercoid of Jlymenolepis imvrina 12 

3. Pedigree of Mr. Smalley's pigeon (W 69) 15 

4. Genealogical Table showing matings and results in mixed strains 

of pigeons 17 

5. Upper, lower, lateral and posterior aspects of skull of male 

Lacerta jacksoni 20. 

6. Lamina supraciliaris and supraorbital bone of Lacerta jacksoni. 

Also showing configuration of nasal process of prsemaxillary 
and outer nasal apertures of L.jacJcsoni, c? , $ > L. miiralis, 

f. typica, S ; L. vivipara, S j ^^^^ L. agilis, c? 23 

7. Upper, lower, lateral and posterior aspects of skull of female 

Lacerta Jacksoni 27 

8. Upper and side views of head of male a'ld female Lacerta 

peloponnesiaca ^^8 

9. Intestinal tract of Ithea americana 52 

10. Intestinal tract of Crypturus tataupa , 53 

11. Intestinal tract of Talegalla lathami 55 

12. Intestinal tract of Ortalis ruficatida - 56 

13. Intestinal tract of Euplocamus nycthemerus, showing condition 

reversed from the normal 57 

14. Intestinal tract of Podargns cuvieri 58 

15. Intestinal tract of Gymnorhina leuconota 59 

16. Intestinal tract of Melanerpes superciliaris 62 

1 7. Intestinal tract of Gecinus viridis 63 

18. Intestinal tract of Grus japonicus , 65 

19. Intestinal tract of Turnix varicc 67 

20. Intestinal tract of Spizaetus hellicosus = . , . 09 

21. Intestinal tract of Cathartes aura 70 

22. Intestinal tract of Asio otus 71 

23. Intestinal tract of Eupodotis aiistralis 74 

24. Intestinal tract of Pluvianus cegyptius 75 

25. Intestinal tract of (Edicnemus scolopax 76 

26. Intestinal tract of Frafercuia arctica 79 



XIX 

Page 

27. Intestinal tract of Tachyhaptcs Jiuviatilis , 81 

28. Intestinal tract of Nestor notabilis 84- 

29. Dissection of a full-grown Axolotl {Amblystoma tiyrhmni), 

showing the skeleton and nerves of the head and three trunk- 
segments 104 

30. Outline of spinal cord and hind brain, with roots of cranial 

nerves and first two spinal nerves, of a full-grown Axulotl . . 105 

31. 32. Partial reconstructions of the hind region of the skull and 

the anterior region of the trunk of an advanced larva of the 

Axolotl 105, 106 

33-35. Partial reconstructions of the left side of the head region of 

a young larva of the Axolotl 10(5, 107 

36. Reconstruction, seen from above, of the hind region of the skull, 

and anterior neural arches of a young larva of the Axolotl . . 107 

37. Left side view of a partial reconstruction of the anterior end of 

the trunk-region of a young larva of the Axolotl 108 

38. 39. Partial reconstructions, seen from the right side, of the 

anterior end of the trunk-region of a young larva of the 
Axolotl 108 

40. Reconstruction, seen from above, of the head and anterior 

trunk-region of a young larva of the Axolotl 109 

41, 42. Reconstructions of the anterior trunk-region, seen from the 

right side, of a young larva of the Axolotl HO 

43. Left side view of a reconstruction of the anterior trunk-region 

of a young larva of the Axolotl Ill 

44. Right side view of a reconstruction of the anterior end of a 

larva of the Axolotl " Ill 

45. Partial reconstruction, seen from above, of the anterior region of 

an embryo of the Axolotl 112 

46. Similar reconstruction of the more dorsal region of the same 

Axolotl embryo 113 

47. Left side view of a reconstruction of the anterior region of an 

embryo Axolotl 113 

48. Partial reconstruction of the anterior region of an embryo 

Axolotl, seen from above 114 

49. Right side view of a reconstruction of an embryo Axolotl .... 115 

50. Diagrams of the segmentation of the metaotic region in 

Amphibia 116 

61. Diagram of the segmentation of the anterior region of an 

Amphibian 116 

62. Head of a Caribou [Rangifer tarandus caribou) with abnormal 

antlers 182 

53. The bladder, duct, and " red bodies " oi Anguilla vulgaris, viewed 

from the side 186 

54. External aspect of an Eel's rete mirabile 187 

55. The construction of an Eel's rete mirabile bipolare geminum . . 188 



XX 

Page 
56, Vein surrouuded by modified acini of the pancreas in Neropkis 

(squorius ..,...,, 199 

67. The construction of the " red "body " of Peristethus cataphractus . 202 

68. Diagram to explain the probable construction of the gas glands 

of Trigla gwnardus and Smarts maurii 207 

69. Diagrammatic transverse sections tbrough the anterior end of 

" red body " in ventral wall of bladder of Cepola ruhescens . . 213 

60. Vacuoles in the kidney cells of Neropkis 225 

61. Microphotograph of transverse section across thin region of gas 

gland of Perca, showing foam-like mass of bubbles on surface. 240 

62. Microphotograph of transverse section across thick region of gas 

gland of Perca, showing extruded gas bubbles on surface. 
The " spent " condition of the cells is also shown , . 240 

63. Skull of the Anoa {Bos dejyressicorms) 250 

64. Front part of the skull of a Buffalo (Bos hubaliis) 251 

65. Front part of skull of the Urus (Bos j^rimigenius) 251 

66. Skull of a Urus in the Anatomical Museum, University of 

Edinburgh 252 

67. Front part of skall of American Bison 253 

68. Front and side views of the skull of a young foetal Ox 253 

69. Front view of a fcetal Ox skull about the fifth month, and front 

part of same skull 255 

70. Front view of Calf's skull at birth, and front part of same skull . 257 

71. Front part of skull of a polled Aberdeen- Angus Ox 258 

72. Skull of a Syrian Ox with vestigial horn-cores and a forehead 

like that of flat-polled Aberdeen-Angus cattle 259 

73. Front part of skull of polled Newstead Ox in which the upper 

end of the premaxilbie bifurcntes 261 

74. Front part of a horned NeAvstead skull in which the premaxillse 

are short and fail to reacli the nasals 262 

76. Front part of skull of a borned Cadzow Ox with notched pre- 

maxillse which fail to reach the nasals 263 

76. Front part of skull of a cross-bred Shorthorn in which the pre- 

raaxillte fail to reach the nasals 263 

77. Skull of polled Newstead Ox of the type, which, witli hoins, are 

sometimes said to belong to Bos frontosus Nilsson 264 

78. Eight half of occiput of the Aberdeen-Angus skull with long 

premaxilhe. Also a section of same 265 

79. Outlines of the occiput and part of the forehead of three Urus 

skulls 266 

80. Front view of skull of the Bison [Bos bkon) 267 

81. Hind part of skull of the Urus represented in text-fig. 66 268 

82. Frontlet and horn-cores of a small Newstead Ox with a 

prominent forehead 269 

83. Occiput of the Urus skull represented in text-figs. 66, 79, and 81. 270 



XXI 

Page 

84. Occiput of the Newstead skull represented in text-fig. 86 .... 271 

85. Occiput and horn-cores of a Urus in which the notches below 

the horn-cores are deeper than in text-fig. 83 272 

86. Front view of the Newstead skull of which the occiput is 

represented in text-fig. 84. Also the temporal fossa of same 
skull 274 

87. Front view of skull of a white "wild" Cadzow Ox from 

Hamilton Park. Also occiput and horn-cores of same 
skull 275 

88. Occiput and horn-cores of an American Bison 276 

89. Front part of the skull of a Zebu [Bos inddcus) in which the 

premaxillsD reach the nasals 278 

90. Front part of the skull of a Zebu in which the premaxillee fail 

to reach the nasals 279 

91. Occiput of the Zebu with the premaxillte reaching the nasals. . 280 

92. Tetraijnatha ferox, var. crossce. Male 301 

93. TJliodon haivkesi. Female 304 

94. A rgoctenus aureus. Female 306 

95. Mynoglenes chiltoni 309 

96. Dolomedes trirlentatus. Female 311 

97. Young Brindled Gnu born in the Society's Gardens 315 

98. Head of young Brindled Gnu , 316 

99. Diagram to explain the tables of different markings in Scila- 

mandra maculosa , . . 325 

ICO, 101. Salamandra maculosa forma typica 329, 332 

102. Salaviandra tnaculosa var. tcsniatu 339 

103. Head of an old Buck of the Mountain Nyala 351 

104. Clasping organs of Arcliceopsylla sinensis S 366 

105. Abdominal segments VII & VIII and receptaculuni seminis of 

Arcliceopsylla sinensis 5 .' 366 

106. Head of Ceratophyllus o'ispus c? 367 

107. Clasping organs of Ceratophyllus crispus (^ 370 

108. Abdomir.al segments VII & VIII, stylet and receptaculuni 

seminis of Ceratophyllus crispus 5 371 

]C9. Abdominal segments VII & VIII and stylet of Ceratophyllus 

dolabris 2 , 371 

1 1 0. Clasping organs of Ceratophyllus dolabris S 373 

111. Abdominal segment VIII and receptaculum seminis of Cerato- 

phyllus crassus $ 375 

1 1 2. Clasping organs of Ceratophyllus mandarinus S 377 

113. Abdominal segments VII & VIII and receptaculum seminis of 

Ceratophyllus mandarinus 5 378 

114. Clasping organs of Ceratophyllus monyoUcus S 379 

115. Abdominal segments VII & VIII aid reteplaculuni seminis of 

Ceratophyllus famulus § 381 



XXll 

Page 

116. Abdominal segments VII & VIII and receptaculum seminis of 

Ceratophyllus euteles 2 383 

117. Ahdominal segments \11 & \III oi Cerafophi/llus phoiopis .. 383 
lis. Clasping organs oi Amphvpsylla casis J 386 

119. Abdominal segments VII & VIII and receptaculum seminis of 

Amphipsylla casis 2 • • ■ 386 

120. Abdominal segments VII & VIII and receptaculum seminis of 

Neopsi/lla compar 5 387 

121. Clasping organs of Neopsylla aliena cJ 390 

122. Abdominal segments VII & VIII and receptaculum seminis of 

Neopsylla aliena 5 390 

123. Head of Stenoponia coelestis § 392 

124. Abdominal segments VII & VIII of Stenoponia ccelestis $ .... 392 

125. Ventral view of anterior part of the body of Megalophrys fece 

partially dissected 394 

126. Sternum of Megalophrys fees from the vential surface 395 

127. The alimentary tract of i^fe;{7a/o/>7«'?/s /6-(B from the middle of 

the stomach to the middle of the colon laid open and divided 
into two portions 398 

128. Gular sac of Megalophrys fecB 401 

129. Ventral musculature of the throat of Megalophrys fece 403 

130. Alimentary tract oi Breviceps gibbosus with the greater part of 

the small intestine laid open 407 

131. Breviceps gibbosus, from the dorsal surface, partly dissected . . 409 

132. An enlarged view of a portion of text-tig. 131, to show absence 

of diverticulum of body-cavity overlying thigh 410 

133. Certain parts in the anatomy of Breviceps sp 411 

134. Spermatheca of PJieretima moiitana containing spermatophores . 414 

135. Longitudinal section through spermatophore of Pheretima 

montana 416 

136. Transverse section through wall of spermatophore of Pheretima 

montana 417 

137. Mesoplodon {Dioplodan) grayi 420 

138. Palatal aspects of cranium and lower jaw of Mesoplodon 

[Dioplodon) grayi , 421 

139. Lateral view of skull of Mesoplodon (Dioplodon) grayi 422 

140. Acheilostoma simpsoni 650 

141. Cylindropharynx brevicauda 552 

142. Cylindropharynx longicauda 553 

143. Trachypharynx nigerice 554 

144. Spiroptera uganda 555 



NEW GENERIC TERMS 



PROPOSED IN THE PEESENT VOLUME (pp. l-5or)). 



Page 
Aclieilostoiua (Vermidea) 549 

Cobboklina (Vermidea) 655 

Corycella (Crustacea) 283 



Page 
Cylindropharynx (Vermidea) ... 551 

Stenoponia (Siphonaptera) 391 

Trachypliarynx (Vermidea) ... 552 



INDEX. 



Abdiitiia 

sphenorhijncha, aOO. 
Aburria 

carunculata, 523. 
Acanthi on 

hrachyura, 14"). 
Acanthodactylns 

hoskianus, 30. 
Accentor 

collaris, 471. 

modularh, 47 1 . 
Accipiter 

melanoleucua, 50-'!. 

nmis, 503. 

pileatus, 503. 
Aceros 

nepalensis, 4S8. 
Acheilostoma, gen. nov., 
649. 

simpsoni, 549, 550. 
Acomys 

cahirensis, 5. 

hunteri. 448. 
Acridotheres 

cristatellus. 481. 

Jiiscus, 481. 

ginginianus. 481. 

onahratfensis, 481. 

^r/s;'/s, 481. 
Acrocephalus 

strepems, 471. 
Acryllium 

vulturinum, 523. 
Add ax 

naso-maculat'Us, 1, 456. 
TEchmophorus 

major, 532. 
jEffialitis 

citronica, 529. 

hiaticula, 529. 
..^luroedus, 86. 
jElurus 

fulgens, 442. 

Proc. Zool. Soc— 19 



11 



.-Epyprymniis 

rufescens, 464. 
Aex 

galericidafa, 512. 

sponsa, 512. 
Agapornis 

cawre, 498. 

imllaria, 498. 

roseicoUis, 498. 
Agelteus 

frontalis, 481. 

phcenicetis, 481. 

rvjwapiUus, 481. 
Aj«ja 

TOsea, 1, 510. 
Akis 

sphtosa, 10. 
Alactaga 

decumana, 449. 

indica, 449. 

jaculus, 12. 
Alauda 

arborea, 483. 

arvensis, 483. 

ccelivnx, 484. 

crassirostris, 484. 

crislata, 484. 

qulqida, 484. 
Alca ■ 

?'orf/«, 532. 
Alcedo 

M?32cfff, 64, 91, 488. 
Alce^ 

ff/ces, 151. 

machlis, 459. 
Algiroides 

a.fricwnus, 30. 

nigropunctatii^, 22. 
Allactaga 

lovgipen, 148. 
Alma, 414. 
Alopex 

lag opus, 135. 
. No. XXXYII. 



Amadina 
hicolor, 475. 
castanotis, 475. 
erythrocephala, 475. 
fasciata, 475. 
fringilloides, 475. 
lathami, 475. 
modesta, 475. 
Amblyrhamphus 

holosericeus, 481. 
Amblystoma, 104, 105, 
115, 118. 
tigrinum, 103. 
Ammoperdix 
honhami 621. 
%i, 521. 
Ampelis 

cedronim, 473. 
garrulus, 473. 
Ainphipsylla, 388. 
raszs, 385, 386. 
(?««, 385, 386, .387. 
Amyntas, 413. 
Anas 

hoscas, 513. 
crisiata, 513. 
ohscjira, 513 
pcecilorhyyicha, 513. 
specularis, 514. 
superciliosa, 614. 
undulata, 514. 
Ancylostomura, 549. 
Anguilla, 183, 191, 192. 
193, 194, 196, 216, 
217, 219, 237. 
vulgaris, 185, 186,191, 
192, 245. 
Anisolabis 

annulipes, 10. 
Anoa 

depi'essicortiis, 454. 
Anodorhynchns 
glaums, 495. 

37 



INDEX. 



Anodorliynclius 

hi/acintJmms, 493. 

Icarl, 405. 
Anourosorex 

squamipes, 16S. 
Ansii" 

alhifrons, .512. 

hj-achi/rhi/nrkus, 512. 

cinereus, 512. 

cyanoides, 612. 

erijihroipus, 513. 

iiidicus, 513. 

^egetum^ 513. 
iliiserana.s 

ssmipalmata, 512. 
Anteliomys, 177. 
AiUhornis 

onelanura, 474. 
Anthracoceros 

coronatus, .488. 

malabaricus, 488. 

nnalayanus, 488. 
Anthropoides 
caruneidata, 527. 
leitcauchen, 83, 527. 
Jeucogeranos, Ty21 . 

paradisea, 4, 53, 527. 
virgo, 527. 
Anthropopitheciis, 121, 

.).5<J. 

caluus, 429. 

gorilla, 429. 

sati/rus, 124, 125. 

troglodytes, 354, 429. 
Aiithus 

arboreus, 472. 

ohscurus, 472. 

pratensis, 472. 

spipoletta, 472. 
Antilocapra 

americana, 458. 
Antilope 

cervicapra, 152, 456. 
Apbobus 

chopi, 481. 
Apodemus 

agrarius, 172, 173 

— ningjKensis, 173. 

— paliidior, 172, 173. 
fergussoni, 172. 
speciosus chevrieri, 172, 
sylvaticus, 148. 

Aprosniictns 

coccmopterits, 498. 

cyanopygius, 498. 
Aptenodytes 

pennanti, 632. 
Apteryx, 55, 59, 88, 91 
93. 

aitstralis, 61, 533. 

haasti, 533. 



Apteryx 

riiantclli, 51. 533. 
owsni, 533. 
Atpiila 

adcdberti, 50l!. 
audax, 502. 
chrysaelus, 502. 
claiiga, 502. 
imperialis, 502. 
ntsvioides, 502. 
verreauxi, 602. 
Ara 

ambigtia, 495. 
ararauna, 83, 84, 85, 

495. 
chloroptera, 495. 
hahni, 495. 
macao, 495. 
maracava. 495. 
miliiaris, 84, 495. 
oiobilis, 495. 
severa, 495. 
Ara.mides 

cayennensis, 525. 
ypiecaha,, 82, 525. 
Aramus 

scolopaccus, 528. 
Araneus 

verrucosus, 297, 299. 
Arboricola 

atrogularis, 520. 
torqucola, 520. 
Arcella, 229. 
Archojopsylla 
eriiiacei, 365. 
sinensis, 365, 366. 
Archibuteo 

lagopus, 501. 
Arctic'tis 

biniurong, 181, 438. 
Arctogale 
leucotis, 438. 
trivirgata, 438. 
Arctoniys 

caudatus, 446. 
himalayanus, 446. 
hodgsoni, 416. 
marmotta, 446. 
monax, 446. 
] Ardea 

aqmni, 608. 
! atoff, 508. 
argala, 49. 
atricollis, 508. 
buh ulcus, 508. 
candidissima, 508. 
cinerea, 508. 
coco/, 508. 
egretta, 508. 
garsetta, 508. 
goliath, 508. 



Ardea 

gularis, 608. 
herodias, 508. 
novcB'hollandia, 608. 
purpurea, 508. 
ralloides, 508. 
sumatrana, 508. 
Ardetta 

involucris, 508. 
minuta, 608. 
Argoctemis, 297. 
aM/-e«.s, 299, 306. 
vitfatiis, 398. 
Argus 

giganteus, 622. 
Art.amus 
fuscus, 473. 
personata, 473. 
sordidus, 473. 
Stvperciliosus, 473. 
Arricanthis 

pulchellus, 124, 148. 
striatus, 124, 148. 
Arvicola 

agrei^tis, 448. 
amphibius, 147, 448. 
pratensis, 448. 

terrestris, 147. 
Asarcornis 

scutulatiis, 4. 
Asio 

brachyotus, 492. 

mexicamis, 492. 

oiJMs, 71, 72, 492. 
Asopia 

farinalis, 10. 
Astur 

approximans, 68, 603. 

novm-hollandicB, 503. 

palumbarius, 68, 503. 

tachiro, 503. 

tibinUi, 503. 
Asturina 

magyiirostris, 502. 

nattereri, 502. 

nitida, 602. 
Ateles 

a/er, 433. 

geoffroyi, 433. 

grisescens, 433. 

hybridus, 433. 

melanochir, 433. 

paniscus, 433. 

rufiventris, 433. 

variegatus, 433. 

vellerosus, 433. 
Athene 

brama, 493. 

noct'ua, T2, 493. 
Atlierina, 183, 204, 206, 

209, 219. 



IXUEX. 



Atherina 


Bibos 


Box, 183, 219. 


hepsetHs, 20o, 216, 247. 


gaurus, 451. 


/)w^«, 205, 216, 247. 


AtLierura 


Bison 


B.-adypus 


africfiiia. 449. 


(tniericanas, 100, 454. 


di'lach/Ius, 132. 


fasctciiJata, 4i9. 


/;Jso«, 154. 


friduc'tylus, 132, 462. 


Atlierunis 


honasus, 154, 454. 


Breviceps, 397, 399. 


macroura, 145. 


Bitis 


sp., 405, 408, 411. 


Atlantoxerus 


nasicornis, 2. 


adspersus, 404. 


gciulus, 149. 


Biziura 


(/ihbosus, 401, 404, 405. 


Aulacodiis 


lobata, 514. 


4U6, 407, 408, 409, 


swindcniianuf. 449. 


Blariua, 166, 167. 


410, 412. 


Aulacorhamphiis 


Blarinella, gen. no v., 1 59, 


macrops, 404. 


sulcadis, ti3. 


166. 


mossambicvs, 404. 




quadraticauda, 166, 


pantheri, 404. 




. 167. 


vnrrucosus, 404. 




(Cryptotis) p«rm, 167. 


Brotogerys 


Babirussa 


Blcistocerus 


chrysopterus, 496. 


al funis, 4G1. 


bezoarticus, 151. 


jugnlaris, 496. 


bahirussa, 140. 


Blennius, 204. 


pyrrho2)terus, 496. 


Balrena 


Bolborbynchus 


(■(V/m, 496. 


hoops, 156. 


' iineolatus, 49(i. 


)'«/, 496. 


muscidus, 156. 


Bos 


tuipara, 496. 


mi/sticetus, 155. 


1 acutifrons, 256, 273, 


virescens, 496. 


phi/sahis, 156. 


' 277,278,279,281. 


Bubal i 3 


Balt^noptera 


i /)?'so», 154, 267. 


boselaphus, 454. 


boopx, 156. 


' honasus, 154. 


caama, 454. 


muscidus, 156, 423. 


bj-achyceros, 249, 254, 


^ra, 454. 


physcdus, 156. 


256, 279, 2.'^0. 


Bubalus, 260, 264. 


Bcilearica, 82. 


hnhalus, 154, 251, 259. 


cequinocticdis, 454. 


pavonina, r>'21. 


chincnsis, 26-1. 


buhalis, 154. 


reqidorum, 527. 


dcpressicnrms, 250, 


huffelus, 4, 454. 


Bali'stes, 183, 215, 219. 


259. 


cff,^er, 454. 


capri.scus, 209, 210, 


frontosus, 249, 264, 


Bubo 


248. 


268, 282. 


ascalaphus, 492. 


Bambusieola 


zMf/era^s 164, 249, 278, 


hevgalensis, 492. 


/v''^'/i'^ 520. 


454. 


bla/dstoni, 492. 


thoracica, 520. 


longifrons, 249, 254, 


capensis, 72. 


Barbastella 


i:56, 262, 279, 280, 


cinerascens, 71, 72. 


darjelingensis, 160. 


281. 


coromandus, 492. 


Bassaricyon 


namadicus, 254, 256, 


lacieus, 492. 


aJ/e?fi, 442. 


264, 272, 273, 277, 


macidosus, 71, 72, 492. 


Bassaris 


281. 


7na gel' aniens, 492. 


astida, 442. 


planifrons, 2.')6, 273. 


maximus, 71, 72, 492. 


Bas.sarisciis, 137. 


primigenius, 249, 250, 


ncpalensis, 493, 


Baza 


25i, 254, 256, 258, 


orientalis, 493. 


lophotes, 504. 


262, 264, 273, 277, 


poensis, 493. 


Benhamia, 418. 


278, 279, 280, 281, 


virginianus, 71 , 72, 


austeni, 414. 


282. 


493. 


Bernicla 


te<res, 154. 249, 454. 


Buceros 


brent a, 513. 


— brachyceros, 254, 


cylindrica, 60. 


canadensis, 513. 


266. 


lunatus, 488. 


hutchinsi, 513. 


— macj-occros, 254. 


rhinoceros, 488. 


lettcopsis, 513. 


— primigenius, 249, 


Bucorvus, 69. 


ruficollis, 513. 


256,258,281. 


abyssinictis, 61, 488. 


Bettongia 


trocJioceros, 250. 


cff/er, 488. 


■ cuniculus, 464. 


Boselaphus 


Bungarus 


gaimardi, 464. 


tragocamelus, 456. 


fasciatus, 4. 


lesueuri, 464. 


Botanrus 


Biiteo 


penicillata, 464. 


lentiginosus, 508. 


alhicaudatvs, 50'. 


Bibos, 260, 264. 


siellaris, 508. 


aiiguralis, 501. 


frontcdis, 454. 


Bothrioneuron, 414. 


hcrealis. 601 . 
37* 



INDEX. 



Buteo 

di'sertontiiifbOl. 
eri/thronotus, 5D1. 
ferox, 501. 
jackal, 601. 
pennsylvardcus. 501 . 
poliosonms, 6U1. 
vulgaris, 501. 
Buteogallus 

nigric&Uis, 502. 
Butorides 

airicapillus, 508. 
ci/antcrus, 508. 
vlrescens, 508. 
Bycanistes 

subcylindricus., 483. 



Cabassous 

unicinctus, 141. 
Cacatua 
aZ6a, 494. 
citrina, 494. 
ducorpsi, 494. 
galerita, 495. 
ffoffini, 495. 
gymnops, 495. 
hamaturopygia , 495. 
leadhcateri, 495. 
moluceensis, 495. 
ophthalmica, 495. 
roseicapilla, 495. 
savguinea, 495. 
sulphurea, 84, 495. 
triton, 495. 
Cdccabis 
chukar, 521. 
melanocephala, 521. 
petrosa, 521. 
rtf/"a, 521. 
saxatilis, 521. 
Caica 

leucogastra, 497. 
melanocephala, 497. 
xanthomtra, 497. 
Cairina 

mos^cliata, 512. 
Calaudrella 
batica, 484. 
brachydactyla, 484. 
Calcai-ius 

lapponicus, 480. 
Galidris 

arenaria, 529. 
Callipepla 

califormca, 521. 
gambelli, 521. 
sqiiamata, 521. 
Calliste 

brasiliemis, 474. 
fastuosa, 474. 



Calliste 
f estiva, 474. 
melanota, 474. 
tricolor, 474. 
Callithrix, 4^3. 
jacchus, 127. 
Callooeplialon 

galeaUim, 84, 494. 
Callorhinus 

iirsinus, 133. 
Cal jcitta 

formosa, 483. 
Calodromus, 57. 
CaicBDas 

nicobarica, 518. 
Calopelia 

puella, 518. 
Calopsittaeus 

novcB-holkmdics, 495. 
CaUiromys 

2)hilander, 143. 
Calyptorhynchus 

banksi, 494. 

funercus, 494. 
Cambridgea 

antifodiana, 297, 300. 
Camelus, 122. 

baciriaoms, 149, 150, 
460. 

dromedarius, 149, 150, 
460. 

glama, 150. 

paces, 150. 
Cancroma 

cochlearia, 508. 
Canis 

alopex, 134. 

antarcticus, 440. 

anthus, 440. 

argentatus, 440. 

aureus, 135, 440. 

azarm, 440. 

bengalensis, 440. 

cancrivorus, 440. 

<?ey-(^o, 440. 

cliama, 440. 

dingo, 440. 

famelicus, 440. 

familiaris, 134. 

fidvicaicdus, 440. 

falvipes, 440. 

fulvus, 440. 

griseus, 3. 

hodophylax, 440. 

hymna, 134. 

jubatus, 440. 

lagopus, 135, 440. 

latercdis, 440. 

latnins, 440. 

leucopus, 440. 

/«/>?«, 134, 440. 



Canis 

magellanicus, 440. 
mesomelas, 440. 
microtis, 440. 

niger, 440. 

iiiloticus, 440. 
pallidus, 440. 
pallipes, 440. 

t^efo.r, 440. 

virqinianus, 440. 

•TO^^jcs, 134, 440. 

(Cuod) primisuus, 
440. 

( — ) rutilans, 440. 

(Nyctereutes) pro- 
cyonoides, 440. 
Capra 

(Bgagrus, 457. 

ammon, 153. 

caucasica, 457. 

cervicapra, 152. 

depressa, 152. 

clorcas, 153. 

gazella, 152. 

grimmia, 153. 

hircus, 152, 457. 

hispanica, 457. 

^7J^f.^■, 152, 457. 

mambrica, 163. 

megaceros, 457. 

niihiana, 457. 

pygmea, 152. 

reversa, 152. 

ruficapra, 152. 

sinaitica, 457. 
Capreolus 

bedfordi, 180. 

caprtBa, 459. 

capreolus, 161. 

pygargus, 459. 
Capriaiulgus 

europcBUS, 487. 
Capromys 

brachyurus, 449. 

^zYorz^cs, 449. 
Caranx, 183, 219. 

trctchurus, 209. 
Cardinalis 

virginianus, 478. 
Carduelis 

elegaiis, 479. 

orientalis, 479. 
Oar lac us 

campestris, 151, 459. 

gymnotis, 459. 

leucurus, 459. 

inacrotis, 459. 

mexicanus, 459. 

ncmorivagus, 459. 

riifus, 459 

virginianus, 459. 



Cariarua 

cristaia, 527. 
Oarphibis 

spinicollis, 510. 
Carpococcyx 

radiatus, 54. 
Carpodacus 

erylhrinus, 479. 

h(e7)iorrhous, 479. 
Carpophaga 

cenea 85, 516. 

coiicinna, 85. 

latrans, 85. 

paulina, 516. 
Oaryoaiys, 175. 
Cassious 

Ji(ematorrhou.s, 480. 

persicus, 480. 
Ca.ssidix 

oryziwra, 60. 
Castor 

canadensis, 447. 

/i^er, 146, 447. 

■mo.ichafus, 146. 
Casuarius 

a.ustraUs, 51. 

hennetti, 634. 

hicarunc'idatus, 534. 

casuarms, 534. 

— ausiralis, 534. 

— beccarii, 534. 

— intensus, 534. 

— salvadori, 534. 

— sclaiori, 534. 

— violicoUis, 534. 
intensus, 87. 
pajnianus, 534. 

— edwardsi, 534. 
philipi, 534. 
pictkollis, 534. 
rothschildi, 87. 
uniapptendiculattts, 

534. 

— aurantiacus, 534. 

— occipitalis, 534. 

— rufotiiictus, 534. 
westermanni, 51, 87. 

Cathai'tes, 61. 

atratus, 505. 

awm, 69, 70, 505. 

uruhitinga, 505. 
Cavia 

flavidens, 450. 

porcell'us, 146. 

s/?m, 450. 
Cebus 

albifrons, 432. 

tf^e^J/ffl, 128, 432. 

azarm, 432. 

capucinus, 128, 432. 

fatuellus, 432. 



Cebiis 
jlavescciis, 432, 
hypoleucKAi, 128, 432. 
lunatus, 432. 
moncwkus, 432, 
vellrrosus, 432. 
versicolor, 432. 
CenteLes 

ecandcdus, 444. 
Centropiis 
phasiaiidos, 490. 
rvfipennis, 490. 
senegalensis, 490. 
Centuriis 

tricolor, 487. 
Cephalophus 
abyssinicus, 2. 
coronat'is, 455. 
dorsalis, .314, 455. 
grim mi. 153, 455. 
riidxwelli, 455. ■ 
monticolor, 455. 
nigrifrons, 455. 
rufiilatti^, 455. 
sylvictdtrix, 455. 
Cepola, 183, 216, 217, 
218, 219. 
ruhescens, 212, 248. 
Ceratodus, 118. 
forsteri, 119. 
Ceratogyiniia 
c/am, 488. 
Ceratophyllus, 388. 
crasaus, 374, 375. 
crispus, 365, 367, 370, 

371. 
c?oZa/;m, 371, 373, 374, 

375. 
eutelcs, 382. 
faimoliis, 380, 381. 
fasciatus, 5, 9, 11. 
'laqomys, 371, 372, 

'373. 
londiniensis, 5. 
mandarinus, 375, 377, 

378. 
mongolicus, 378. 
phceopis, 383, 384. 
tcsquorum, 379, 380. 
Cerchneis 

ardesiacus, 3. 
Cercocebus 
athiops, 431. 
cdhigeiia, 431. 
aterrimus, 431. 
collaris, 431. 
chrysogaster, 431. 
Juliginosus, 431. 
hagenhecki, 431. 
Oercoleptes 

caitdii'olvnlus, 442. 



Oercopithecus 

cetUops, 128, 431. 

alhogidaris, 430. 

a&canias, 430, 

barhatus, 126. 

callitriclms, 431. 

campbelli, 430. 

cephus, 127, 430. 

cynos'urus. 431. 

tZea?«a, 127, 430. 

arythrotis, 430. 

grayi, 430. 

grismviridls, 12S, 
431. 

hdnatus, 430. 

lalandii, 431. 

leucampyx, 430. ' 

rhocsti, 430. 

■Martini, 430. 

mona, 430. 

neglectus, 430. 

nictitans, 430, 

■patan, 431. 

petaurista, 430. 

pygerythrv.s, 431. 

pyrrhonotus, 431. 

sabceus, 431. 

sohmidii, 430. 

stairsi, 430. 

talapoin, 431. 

tantalus, 431. 
Cereopsis, 513. 

novcB-hollandim, 512. 
Ceriornis 

6^?/i;^2, 522. 

caftoili, 522. 

melanocephahc, .522. 

safyra, 522. 

temmincJci, 522. 
Cerodon 

rupestris, 450. 
Cervicapi'a 

bohor, 455. 

isahcUina, 455. 
Cervulus 

crinifrons, 4.58. 

lacrymans, 458. 

munlja.c, 458. 

reevesi, 458. 
Cerviis 
a/ccs, 151. 
a/Arcc?;-, 459. 
aristotelis, 459. 
f7.r2s, 459. 
bezoarticus, 151. 
camelopardalis, 150. 
canad.ensis, 100, 
I 458. 

j dama, 151. 

davidio.nus, 459. 
I diivaucelli, 459. 



INDEX. 



Cervus ] 

elaphus, 150, 151, 

458. 
eldi, 459. I 

hippelafjlms, 459. j 

Jcashmiriatma, 4ri8. | 

huJdi, 459. 
luehdorfi', 458. 
onantchuricus, 459. 
maral, 458. 
moluccensis, 459. 
philippinus. 459. 
porcinus, 459. 
schomburgki, 459. 
s/yto., 459. 
faevanus, 459. 
tarandus, 151. 
Chalcopelia 

chalcospilos, 518. 
Cbalcophaps 

chrysocMora, 518. 
indica, 518. 
Chalcopsittacus 

scintilla^us, 494. 
Cbamtepelia 
jpasserina, 518. 
tcdpacoti, 518- 
Charadrius 

plnvicdts, 529. 
Charax, 208, 215, 219. 
Chasmorhynchus 
nivetis, 484. 
nudicoUis, 484. 
Chaulelasmua 

streperiis, 514. 
Chaiina 

cn'istata, 511. 
derhiana, 511. 
Chen 

ccendescens, 512. 
hyperboreii^, 512. 
nivalis, 512. 
Chenalopex 
(sgyptiaca, 613. 
juhatu^, 2, 513. 
Chenonetta 

jid)ata, 513. 
Cliera 

progne, 476. 
Chibia 

hottentotta, 473. 
Chinchilla 

lanigera, 460. 
Chionis 
afta, 529. 
minor, 629. 
Chivogaleus 
coquereli, 434. 
ot«7m', 434. 
Chiromys 

madagasca7'icnsis, 434. 



Ghlauiydodera 

maculata, 482. 
Cliloephaga 

onagellanica, 513. 
metanoptera, 513. 
poliocephala, 513. 
riibidiccps, 513. 
Ohlorophonia 
sj-isa, 474. 
■viridis, 474. 
Ohloiopsis 
cmrifroris, 472. 
hardwickii, 472. 
Cbodsigoa 

herezowski, 166. 
hypsibia, 166. 
salenskii, 166. 
amithii, 166. 
Cboloepus 

didaciylus, 132. 
Cbolopus 

didactylus, 462. 
hoffmanni, 462. 
Cbrysochloris 

asiatica, 142. 
Chrysomitris 
harhatub, 479. 
spinoides, 479. 
spinus, 479. 
("m^fis, 479. 
yarrelli, 479. 
Cbrysophrys, 215. 
Cbrysothris 

sciurea, 432. 
Cbrysotis 
(Estiva, 496. 
agilis, 496. 
albifrons, 496. 
amazonica, 496. 
avg^ista, 496. 
auripalliata, 496. 
autuinnalis, 496. 
bodvni, 496. 
bouqueti, 496. 
brcmliensis, 496. 
dufresniaim, 496. 
caymanensis, 496. 
collaria, 496, 
farinosa, 496. 
V'fSiln'a, 496. 
/«sfH 496. 
gnatemalcs, 496. 
! guildi7igi, 496. 
inornatus, 84. 
I leucocephala, 497. 

Icvaillanti, 497. 
1 . mercenaria, 497. 
ochrocephala, 497. 
ocJiroptera, 497. 
panamanensis. 497. 
prairii. 497. 



Obrysotis 

rhodocorytha , 497. 
salvini, 497. 
ventralis, 497. 
versicolor, 497. 
vinacea, 497. 
viridigena, 497. 
^■^"«a^fl, 497. 
xantholora, 497. 
Chunga, 85. 

hurmeisteri , 62, 73, 
527. 
Ciccaba, 72. 
Cicblherminia 

deiisirosiris, 471. 
Ciconia 
«i6a, 509. 
boyciana, 509. 
nigra, 609. 
Cinclu.s 

aquaticus, 471. 
Oircaetus 
gallictis, 503. 
pedoralis, 4. 
Circus 

(sruginosus, .'lOl. 
cincraceus, 501. 
cyaneus, 501. 
gouldi, 501. 
macrunis, 601. 
maurus, 601. 
Gissa 

venatoria, 483. 
Cisaopis 

leveriana, 475. 
Citelhis 

mongolicus, 379, 380. 
Cittocincla 

macrura, 471. 
Clangula 

glaucion, 514. 
Clepsine, 419. 
Cobboldia, 555. 
Cobboldina, gen. noi 
555. 
vivipara, 555. 
Cobus 

ellipsiprymnus, 465. 
uncttiosus, 455. 
Coccothraustes 
melanurus, 478. 
personatus, 478. 
vulgaris, 478. 
Oochoa 

viridis, 473. 
Coelogenys 

7?«ca, 450. 
Coendou 

prcfiensilis, 145. 
Ccereba 
cyanea, 474. 



Colaptes 

anratus, 487. 
Coliopassei" 

maerurus, 476. 
Col i us 

capensis, 487. 

ra.'^tanofus, 487. 

erythromclon, 487. 

niqricolliii, 487. 
Co'.dbus, 12(i. 

vellerosms, 430. 
Coiugo 

■pliilijypinen^i^, 1 oO. 
OuliiiuBa 

alhigularis, 51(). 

draitcana, hlQ. 

hollii. 516. 

carihhcBci, 516. 

eversmaiiui, 516. 

guinea, 51(5. 

gt/mv.ophthalma, 2, 4, 
516. 

i.nornata, 51<\ 

laurivora, 517. 

lencocephala, 517. 

leuconofa, 517. 

maculosa, 517, 

ffw^is, 517. 

■palumbus, 517. 

picazuro, 517. 

plttmhea, 517. 

rujiiia, 517. 

speciosa, 517- 

squamosa, 517. 
Colmiibula 

pici'i, 517. 
Colymbiis 

arcticus, 532. 

glacialis, 532. 

septenirionalis, b'i'2, 
Condylura 

criniafus, 1 43. 
Conepatus 

mapurito, 441. 
Connochistes 

^«M, 454. 

iaurhuis, 181. 
Conuropsis 

carolinensis, 496. 
Conurus 

aciiticaudahis, 495. 

(Sruginosus, 495. 

atireus, 495. 

auricapillus, 495. 

a^'^Jec, 495. 

cactorum, 495. 

canicularis, 495. 

chlorofterus, 495. 

guaroiiha, 495. 

hmmorrhous, 495. ■ 



Conurus 

holochlorus, 495. 

jeiidagi, 495. 

tcucoiis, 85. 

iiandait, 495. 

ocularis, 495. 

ruhritorques, 49G. 

rubrolari'ii/us, 49G. 

sdlslilialis, 496. 

waglp.ri, 49(i. 
Copsychu.s 

mularls, 471. 
Corycias 

c#«e.s, 489. 

garrultis, 489. 

'//(svius, 4. 
Coracoijsis 

harkh/i, 497. 

nigra, 497. 

i^«srt, 497. 
Corcorax 

melauorhamphus, 483. 
Corcgonus 

acroiiiuf, 228. 
Coris, 183,217,219. 

guifredi, 208. 

>/w, 208, 209, 214, 
216, 247. 

vukjari>i, 208. 
Corvina, 183, 215. 

«<V''«- -0^*' -l*^*' -16. 
^217, 248. 
Corvultur 

alhicoli'is, 482. 
Cor \ us 

aniericumos, 482. 

australU, 482. 

ca pell aims, 60, 482. 

corax, 482. 

comix, 482. 

corone, 482. 

cubiiinatus, 482. 

dailricus, 482. 

frugilegus, 482. 

monedula, 482. 

scapulatus, 482. 

splendciis. 482. 
ConcEeus, 282, 283. 

africccnus, 288, 294. 

rt^zYLs 290. 

a^ai-ecs, 287, 288, 289. 

amazonicus, 288, 292, 
293, 294. 

andrewsi, 294, 295, 
296. 

avglicus, 289, 291, 
294. 

adaticus, 288, 294. 

(•«/«(«, 287, 290, 296. 

crassiusculus, 290. 



CoryciEus 

rf««(E, 287, 290. 
cZ?<62«s, 292, 296. 
elonqatus, 287, 238, 

289. 
crgthrceus, 288, 292, 

293, 294. 
yto(;c;<.s, 287, 288, 289. 
furcifer, 287, 288. 
gibbidus, 284, 285, 
gracilicarida.tus, 287, 

290, 296. 
huxleyi, 288. 
feifMs, 287. 

kz(z;;/s, 287, 238, 294. 
limhatus, 287, 289. 
lonqistylis, 287, 288. 
lulibocki, 288, 294. 
mcgalops, 284, 285. 
minimus, 288, 294. 
minutus, 287. 
murrayi, 294, 296. 
obtusus, 287, 289, 29l), 

291, 296. 

oiw//s, 287, 290, 291. 
■pacificus, '2S1. 
■pellucidus, 284, 285. 
robuslus, 287, 289. 
speciosus, 282, 283, 

287, 289. 
/■c^/Kfs, 288, 291, 292, 

294, 296. 
varius, 288. 
venustus, 287, 289, 294. 
vitreus, 287. 

Corycelia, gen. iiov., 

283. 
sp., 237, 296. 
/«-cy/s, 284, 285, 296. 
carinaia, 283, 284, 

287. 
concinna, 283, 284, 

286, 287, 296. 
cicrta, 284, 286, 296. 
^ii^M/rt, 283, 284, 285, 

286, 287. 
gracilis, 283, 287. 
longicatidis, 283, 284, 

285. 
pellucidus, 283, 284. 
rostrata, 283, 284. 
speciosus, 283. 
tenuicauda, 283. 
Corypbospingus 
cristatus, 478. 
pileatus, 478. 
Coscoroba 

Candida, 512. 
Cot-inga 
cincta 484 



xxxii 


INDEX. 




Cotuniix 


Cross opt i Ion 


Cygnus 


chinensis, 63, 520. 


mantchurivma, 521. 


musicus, 512. 


conununis, 520. 


tibetanum, 521. 


nigricoUis, 612. 


coromandelica , 520. 


Crotophaga 


ofor, 512. 


histrionica, 520. 


a«i, 490. 


Cylicostomum, 551. 


pectoralis, 520. 


Cryptobranehus, 117. 


Cyliiidropharynx, gen 


Cr.icrjcus 


Cryptoisrocta 


nov., 551. 


destructor, 483. 


/e/-oa-, 438. 


brevicauda, 551, 552. 


pieatus, 483. 


Cryptotis, 167. 


lonqioauda, .551, 553. 


Cranorrbinus 


Orypturus, 93. 


Cynaslurus, 135. 


corrugatus, 488. 


noctivagus, 533. 


jiibatiis, 4, 438. 


Crax, 67. 


obsoletus, 533 


Cynictis 


a^6eri!j, 523. 


tataupa, 52, 53, 54, 


penicillata, 439. 


alector, 523. 


633. 


Cynocephalus 


carunculata, 55, 523. 


undulatus, 633. 


amibis, 432. 


daubentoni, 623. 


Ctenocephalus 


babouin, 432. 


globicera, 56, .523. 


ca?i/s, 5. 


doguera, 432. 


globidosa, 523. 


/efe, 5. 


hamadrycts, 432. 


hecki, 56. 


Ctenodactylus 


ibeanus, 432. 


incommoda, 523. 


gundi, 449. 


leucophwus, 432. 


sclateri, 523. 


Ctenophthalmus, 388. 


mormon, 432. 


Crex 


Otenopsylla 


neumanni, 432. 


pratensis, 525. 


mayardi, 5. 


papio, 432. 


Cricetomys 


Ciiculug 


po?'carms, 432. 


gambianus, 448. 


canorus, 490. 


sphinx, 432. 


Crieetuliis 


Ourieu.s 


t'otojs, 124, 129. 


a.ndersoni, 173. 


aterrimus, 481. 


Cynomys 


Cricetus 


Cyanecula 


ludovicianus, 446. 


cricetus, 147. 


suecica, 471. 


Cynonyeteris 


fruinentarius, 448. 


Cyanocitta 


coUaris, 444. 


Crithagra 


coronata, 483. 


Cynopithecus 


(dbogularis, 479. 


cristata, 483. 


niger, 432. 


butyracea, 479. 


melanocyanea, 483. 


Cynopterus 


chrysopyga, 479. 


yucatanica, 483. 


marginatus, 444. 


musicus, 479. 


Cyan oco rax 


Cypiinus 


sidphurata, 479. 


cyanomelas, 483. 


m-r/j2o, 228. 


Crocidura 


cyanopogon, 483. 


Cypselus, 60. 


attenuata, 168. 


luxuosus, 483. 


Cystophora 


ccsridescens, 444. 


pileatus, 483. 


cristata, 443. 


Crocodilus, 87. 


Cyanolyseus 




Crocopus 


patagonus, 496. 




chlorngaster, 516. 


Cyanopolius 




plianicopierus, 516. 


cooA'i, 483. 


Dacelo 


Crocuta, 93. 


cyanus, 483. 


cervina, 488. 


capensh, 96. 


Cjanopsittacus 


qigantea, 488. 


crocuta, 95. 


sp/a-z, 495. 


/eac-/z«, 488. 


gariepensis, 95. 


Cyanorhamphus 


Dacnis 


ger^ninaiis, 95, 99. 


auriccps, 499. 


Ci/ana, 474. 


Jcibovotensis, 95, 98. 


malherbi, 499. 


Dafila 


leonticwi, 95, 97, 98. 


nov(B-zealandia, 499. 


f?c?«i'«, 514. 


noltei, 95. 


saisseti, 499. 


spbiicauda, 514. 


vyascB, 99. 


unicolor, 499. 


Daina 


panganensis. 95, 93. 


Cyanospiza 


mesopotamica, 4.59. 


rufopicta, 97. 


cms, 477. 


vv.lgaris, 459. 


thierryi, 95. 


cyanea, 477. 


Danialiscus 


thomasi, 98, 99. 


Cyclopes 


albifrons, 454. 


iogoetisis, 95. 99. 


didactyla, 132. 


pygargus, 454. 


n-issmanni, 95, 96, 97. 


Cygnus 


Dasyproeta 


Crossarchus 


atratus, 612. 


acouchy, 450. 


fasciatus, 1, 439. 


heu'icki, 512. 


d/^F^.?"?", 450. 


obsciirus, 439. 


buccinator, 512. 


azarm, 450. ■ 



xxxui 



Dasyprocta 
cristata, 450. 
isthmica, 450. 
leporinus, 146. 
mexicana, 450. 
'prymnoloplia, 450. 
■punctata, 450. 
Dasypus, 121, 122, 124. 
mill lit us, 462. 
novcmcinctus, 141 , 

142. 
quadricinetus, 141. 
sejitemcinctt/s, 123, 

142. 
sexcinctus, 141, 462. 
triciiictiis, 141. 
unici7ictus, 141. 
villostis, 462. 
Dasyurus 

onaculatiis, 466. 
viverrinus, 466. 
Eaulias 

Inscinia, 471. 
Delpliiniis 
ddjihis, 158. 
wca, 158. 
phoccBoia, 158. 
Dendraspis 
viridis, 3. 
Dendrocitta 

Irimalayensis, 483. 
sinensis, 483. 
vagabunda, 483. 
Dendrocopus 
major, 487. 
Deudrocygna 
arborea, 513. 
arcuata, 513. 
autumnalis, 513. 
discolor, 54. 
eytoni, 513. 
./m^i;«, 513. 
javanica, 513. 
major, 513. 
viduata, 613. 
Deiidrolagus 
bcnnetti, 464. 
■inustus, 464. 
Deroptyus 

accipitrinus, 497. 
Desis 

marina, 298, 299. 
Desniana 

moschatus, 146. 
Dichoceros 

bicornis, 488. 
Dicofcyles 
labiatus, 461 . 
• tajacu, 461. 
Didelphys 
azara, 466. 



Didelphys 

cancrivorus, 466. 

cinereus, 466. 

crassicaudata, 466. 

dorsigera, 144. 

lanigera, 466. 

marsnpialis, 143, 
144. 

murina, 144, 466. 

nudicaudata, 466. 

opossum, 143, 466. 

philander, 143, 466. 

virginiana, 466. 
Didunculus 

strigirostris, 518. 
Dignus 

soiuerbyi, 388. 
Diloplius 

carunculatus, 482. 
DioplodoTi, 422. 
Di])hyllodes 

hunsteini, 2, 60. 
Dipodillus 

campestris, 449. 

simoni, 449. 
Dipus 

cegyjitius, 449. 

hirtipes, 449. 
Dipylidium 

caninum, 12. 
Discoglossiis 

pictiis, 408. 
Dissura 

episcopus, 509. 

onaguari, 509. 
Diiica 

grisea, 477. 
Dolichonyx 

oryzivora, 480. 
Dolichotis 

patachonica, 450. 
Dolomedes, 313. 

tridentatus, 300, 311. 
Donacola 

castaneo-thorax, 476. 

pecturalis, 476. 
Dorcelapbus 

americanus, 314. 
— macruriis, 182. 
Dorcopsis 

liictuostis, 464. 
miilleri, 464. 
Drances 

huttoni, 313. 

frippi, 313. 

Drepanoplectes 

jac/tsoni, 4. 
Dromaeus, 90. 

novcB-hollandice, 534. 
Dromicia 
««7m, 464. 



Uryotriorchis 

spectabilis, 503. 
Dymecodon, 163. 

Echidna 

hystrix, 467. 
Eclectus 

cardinalis, 497. 
pecioralis, 85, 498. 
rorafus, 498. 
westermani, 498. 
Ectopistes 

migratorius, 517- 
Eimei'ia 

(Coccidiuni) avium, 46. 
Elanus 

cmndeus, 504. 
Elaphodus 

rnichianus, 458. 
Elephas 

africanus, 452. 
antiqims, 256. 
indicus, 131, 452. 
maximus, 124, 131. 
Emberiza 
aureola, 480. 
chrysophrys, 480. 
cm, 480. 
ciopsis, 480. 
czr^MS, 480. 
citrinclla, 480. 
hortulana, 480. 
luteola, 480. 
melanocephala, 480. 
miliaria, 480. 
rw^jYa, 480. 
schceniclus, 480. 
striolata, 480. 
Entoniyza 

cyanotis, 474. 
Eos 

riciniata, 494. 
reticulata, 494. 
rubra, 494. 
wallacei, 494. 
Eothenomys, 175. 
Epimys 

andersoni, 171. 
confucianus, 170, 171. 
— luiicolor, 169. 
edwardsi, 171. 
excelsior, 170, 171. 
jerdoni, 170. 
/ff/?5r, 170. 
itorvegicus, 170. 
rattus, 147. 
Equus 

asinua, 155. 
burchelli, 453. 
caballus, 1.54. 
grevyi, 453. 



INDEX. 



Equus 

hemionus, 453. 

— indicus, 453. 

hemippus, 453. 

onager, 453. 

quayga, 453. 

somalicus, 453. 

tceniopus, 453. 

^eSra, 155, 453. 
Eremias 

arquia, 24. 
Eretbizon 

dcvsa^a, 145. 
Erinaceus, 162. 

alhiventris, 444. 

algirus, 444. 

auritus, 444. 

collaris, 444. 

europcsus, 142, 444 

frontalis, 444. 

onicropus, 444. 

miodon, 3(55. 
Erithaeus 

ruhenda, 471. 
Erithizon 

dorsatus, 449. 
Erythrospiza 

githaginea, 479. 
Eli tlir lira 

prasina, 476. 

fsUtacea, 476. 
Esox, 189, 217, 219. 

focms, 190, 215, 243. 
Esti-elda 

amadava, 475. 

fieWrt, 475. 

bichenovii, 475. 

ccsrulescens, 476. 

cinerea, 475. 

cyanogastra, 475. 

dufresnii, 475. 

formosa, 475. 

granatina, 475. 

onelpoda, 475. 

jyhoinicotis, 475. 

phaeton. Alb. 

rubriventris, 475. 

squamifrons, 475. 

subflava, 475. 

temporalis, 475. 
Eudocimus 
«Ztes, 510. 
longirostris, 510. 
ruber, 510. 
Eudromias 

inorinellus, 529. 
Eudynamis 

orientalis, 65, 66, 490. 
ta.itensis, 490. 
Eudyptes 

antipodum, 532. 



Eudyptes 

chrysocome, 532. 

pachyrhynchus, 532. 

sclateri, 532. 
Eii])lionia, 65. 

cJdorotica, 474. 

Jiavifrons, 474. 

laniirostris, 474. 

nigricollis, 474. 

pectoralis. 474. 

sclateri, 474. 

violacea, 60, 474. 
Euphractus, 121, 124. 

sexcinctus, 141. 
Eiiplectes 

«/er, 476. 

capensis, 476. 

jlammicc'2:)s, 476. 

nigriventris, 476. 

ov7/.r, 476. 
Euplooamus 

albo-cristatus, 522. 

aiidersoni, 522. 

erytlirnphtJialmus, 522. 

horsfieldi, 522. 

lineatus, 622. 

melaiLotis, 522. 

nobilis, 622. 

mictliemerus, 57, 58, 
■" 80, 622. 

prmlatus, .522. 

swinhoii, 522. 

vieilloii, 522. 
Eupodotis 

arabs, 626. 

australis, 73, 74, 
526. 

rnjra, 526. 

denhami, 526. 

/l-or2, 4, 526. 

riificoUis, 526. 
Eupsychortyx 

cristatus, 521. 

leylandii, 521. 

sonnini, 57. 
Eiirypyga 

;?e/w.s, 628. 
Euspiza 

americana, 480. 
Exoccetns 
volitans, 228. 



Falco 

(ssa.lon, 503. 
biarmicus, 503. 
concolor, 603. 
eleonorce, 503. 
feldcggi, 69, 503. 
fusco-cisndescens, 503. 



Falco 

lanarius, 503. 
melanogenys, 603. 
peregrinus, 68, 603. 
punicus, 503. 
sacer, 503. 
sidibideo, 503. 
Feiis 

bengalensis, 436. 
cajfm, 437. 
canadensis, 636. 
caracal, 436. 
catus, 136, 436. 
o/ia?fs, 436. 
chrysothrix, 436. 
concolor, 436. 
eyra, 437. 
geofroii, 437. 
fco," 135, 314, 436. 
^^wo^, 136, 436. 
— • isabellinvs, 181. 
maniculata, 437. 
nebulosa, 436. 
owcff., 136, 436. 
pardalis. 136,437. 
2Mrdus, 135, 436. 
passerum, 437. 
planiceps, 436. 
rid)iginosa, 436. 
r/i;/«, 436. 
serval, 436. 

scrvalina, 436. 

temmincJci, 436. 

tigrina, 437. 

fz^rzs, 135, 436. 

uncia, 436. 

viverrina, 181, 436. 

i/agua,rundi, 437. 
Fiber 

zibethicus, 448. 
Fierasfer. 214, 219, 243. 

acifs, 210. 
Foudia 

eryfhrops, 477. 

madagascariensis, 477. 
Francolinu.s 

bicalcaratus, 520. 

cdpensis, 520. 

clappertoni, 520. 

cog'?*/, 520. 

granti, 620. 

gularis, 620. 

/?;?V>(-z, 520. 

levaillanti, 520. 

natalensis, 520. 

pictus, 520. 

ponticerianus, 520. 

vulgaris, 5l'(). 
Fratercula, 59. 

flrc/fm, 78, 79, 80, 
532. 



IXDEX. 



Fregata, 89, 92. 

aquila, 83, 606. 
Fringilla 

Calebs, G, 479. 

kawarahiha, 479.] 

linaria, 479. 

inontifringilla, 6, 479. 

spodiogcnia, 479. 

teydea, 479. 

tintillon, 479. 
Fringillaria 

Sahara, 480. 
Fulica 

ardesiaca, 526. 

ai!ra, 526. 

leitcoptera, 626. 
Fuligula 

6ff.en, 514. 

cristata, 514. 

ferina, 514. 

ferruginea, 514. 

mania, 514. 

rufina, 514. 
Fulinarus 

glacial is, 531. 
Funisciiirus 

leucostigma, 2. 
Fiircifer 

chilensis, 459. 
Furnarius 

rufus, 484. 



GacUis, 183, 212, 215, 
219, 237. 

morrhua, 211, 248. 
Galago 

aWewz, 434. 

crassicaudata, 434. 

garnetti, 434. 

maholi, 434. 

monteiri, 434. 
Galeopithecus, 129. 

■philippincnsis, 130. 
Galeoseoptes 

carolinemls, 471. 
Galictis 

harhara, 441. 

vittata, 441. 
Galidia 

elegans, '439. 
Galliei'ax 

chlorochlamys, 491. 

porphyreolopha, 491. 
Gallinago 

ccelestis, 529. 
Gallinula 

chloropus, 526. 

galeata, 526. 



Gallinula 

vesiotis, 526. 

phceniciira, 526. 

■pyrrhorhoa, 526. 
Galloperdix 

lunulata, 620. 

spadicea, 620. 

eeylonensis, 5-0. 
Gallus 

banJciva, 522. 

sonnerati, 522. 

stanleyi, 522. 

variiis, 522. 
Garrulax 

cdhogularis, 472. 

chinensis, 472. 

leucolophns, 472. 

pcctoralis, 472. 

perspicillat'us, 472. 

'picticollis, 472. 
Garrulus 

glandarius, 483. 

lanceolaius, 483. 
Gasterosteus, 183, 202, 
219, 235. 

aculeat'us, 204. 

spinachia, 201,219. 
Gastropholis 

vittata, 30. 
Gazella 

sp., 180. 

aoxibica, 455. 

bennetti, 455. 

cuvieri, 455. 

dorcas, 153, 455. 

eucliore, 455. 

loderi, 456. 

marica, 456. 

?MoAr, 456. 

muscatensis, 456. 

o'uficollis, 1, 2. 

rufifrons, 3, 456. 

SGemmerringi, 456. 

speJiii, 456. 

suhgutiurosa, 456. 
Geciuus 

t;fmfe, 63, 64^ 487. 
Gelochelidon 

cmglica, 530. 
Genetta 

dongolava, 1. 

jelina, 438. 

genetta, 137. 

pardina, 438. 

senegalensis, 438. 

tigrina, 438. 

tndgaris, 438. 
Geociclija 

ciirina, 470. 

ivardi, 470. 



Geococcyx 

californianus, 65. 
Geopelia 

ciineata, 517- 

humeralis,, 517. 

maugcei, 617. 

striata, 617. 

tranquil la, 517. 
Geophaps 

jdumifera, 518. 

scripta, 518. 
Geop.sittacus 

occidentalis, 499. 
Georychus 

cit'pensis, 448. 
Geotrygon 

cristata, 518. 

montana, 518. 

mystacea, 518. 
Geranoaetus 

inelanoleticus, 68, 
602. 
Gerbillus 

(Bgyptius, 448. 

campestris, 448. 

indicus, 448. 

longifrons, 448. 

pygargus, 448. 

p)yramidum, 448. 

robiistus, 448. 

s^f«6'i, 448. 
Gii-affa, 458. 

camelopardalis, 150. 
Gii-ardinus, 206. 
Glareola 

ocularis, 528. 

pratincola, 529. 
Glaucidium 

passerinum, 493. 

phalanoides, 493. 
Giobicera 

aurorm, 516. 

pacifica, 516. 

rufigula, 516. 
Glossopsittacus 

concinnus, 494. 
Gobius, 183, 196, 204, 
216, 218, 219, 235. 

mjsfiJo, 196, 203. 

micropus, 41, 42, 43, 
44. 

minuius, 40, 41,42,43, 
44, 195, 196, 197. 
219, 239, 246. 

'/«;$^er. 195, 196, 197, 
198, 200, 202, 203, 
219, 246. 

pagunellus, 195, 196, 
197, 198, 210, 214, 
237. 



XX XV I 



INDEX. 



Goura 

coronata, 518. 

victor ice, 518. 
Gracula 

intermedia, 482. 

javauensis, 482. 

religiosa, 60, 482. 
Gr icupica 

nigricoUis, 482. 
Grallina 

australis, 472. 
Grammatoptlla 

striata, 473. 
Gray i a 

sim/thii, 181. 
Grus 

americana, 527. 

antigone, 527. 

austral asiana, 527. 

canadensis, 527. 

communis, 527. 

ja.ponensis, 527. 

japonicus, 64, 65, 
82. 

monachus, 527. 

yiV^o, 82. 
Gubernatrix 

cristo-tellus, 477. 
Guira 

piririgua, 490. 
Guiraca 

ccBTulea, 478. 

cyanea, 478. 

■parellina, 478. 
Giilo 

j9?<^o, 138. 

'luscus, 139, 441. 
Gnttera 

cristata, 522. 

edouardi, 522. 

pucherani, 523. 
Gyninomystax 

■niclanicterus, 4. 
Gymnorhina 

leuconota, 59, 60, 
483. 

!!/6/ce«, 413, 485. 
Gymnura, 162. 
Gypaetus 

harhatus, 505. 
Gypagus, 61. 

V'^)a, 69, 70, 505. 
Gypohieras 

angolensis, 505. 
Gyps 

bengale)isis, 504. 

fulfils, 504. 

/to*i, 504. 

rtteppelli, 504. 



HaBinatopns 

wi^er, 529. 

ostralegus, 529. 
Halcyon 

chtoris, 488. 

sanctus, 64, 91, 488. 

smyrnensis, 488. 

vagans, 64, 488. 
Haliaetus 

alhicillus, 502. 

leucocepJialus, 502. 

leiicogaster, 502. 

leucoryphus, 502. 

vocifer, 502. 
Haliaster 

indus, 502. 

intermedi'us, 502. 
Halichosrus 

gryphus, 443. 
Hapale 

chrysoleucus, 433. 

jaceJius, 433. 

melaniira, 433. 

penicUlata, 433. 

pygmcea, 433. 
Hapalemur 

griseus, 434. 

simus, 434. 
Haplopelia 

larvata, 518. 
Harporhynclius 

rufiis, 471. 
Harpyhaliaetus 

coronatus, 66, 502. 
Hedymeles 

liuloviciamis, 478. 
Helictis 

suhaurantiaca, 441. 
Helotarsus 

ecaudaius, 502. 
Hemicolea 

rogenhoferi, 297, 293. 
Heiiiidactylns 

fasciatus, 3. 
Hemiderma 

brevicauda, 130. 

perspicillatus, 130. 
Ilemigalea 

hardwickii, 438. 
Heiiiitragus 

jemlaicus, 457. 
Hemisos 

fiamla, 472. 
Henieognatbu.s 

leptorhynchus, 496. 
Herpestes 

alhicauda, 439. 

auro-punctatus, 439. 

fulvescens, 439. 

galera, 439. 

gracilis, 439. 



1.55, 



Herpestes 

griseus, 439. 

ichneumon, 439. 

pulverulentus, 439- 

smithi, 489. 
Herpetotheres 

cachinnans, 504. 
rietei'byphantes 

melanugaster , 477. 
Heterolooba 

gouldi, 483. 
Hieracidea 

berigora, 504. 
Hierofalco 

candicans, 503. 

iskaidus, 604. 
Hiinantopus 

brasiliensis, 529. 

■nigricoUis, 529. 
Hippocampus, 208. 
Hippopotairms 

amphibiics, 
461. 

terrestris, 155. 
Hippotr.agiis 

equinua, 456. 

■niger, 456. 
Hiruudo 

rustica, 473. 
Holaspis 

guentheri, 30. 
Homo 

sajDiens, 125. 

troglodytes, 125, 
Houbtira 

macqueeni, 52, 
526. 

undidata, 526. 
Ilysena 

brunncBa., 439. 

cnpensis, 95. 

crocuta, 93, 
439. 

cuvieri, 9.5. 

kycena, 134. 

•)•«/'«, 95. 

.s-^;'vM/«, 134, 439. 
llydrocboerus 

capybara, 450. 
Hydromys 

chrysogaster, 448. 
Hydropotes 

ill cr mis, 459. 
Hydrornis 

a^fewj, 82, 525. 
Hylobates 

agilis, 353, 365, 
3.57,429. 

— martini, 355. 

— pileata, 355. 
gabricllfP, 355. 



94, 



35(') 



XXXVll 



Hjlobates 

hainanus, 353, 354, 

355, 35(1, 357, 429. 
hoolocJc, 355, 356, 357, 

429. 
lar, 355, 357, 429. 
leuciscus, 353, 354, 355, 

356, 357, 429. 
leucogenys, 355, 429. 
mulleri, 355, 429. 
pileatus, 429. 
syndactylus, 429. 

Hylomjs, 162. 
liymenoltBinus 

■malacorhynchus, 514. 
Hyinenolepis 

contractu, IS. 

dimlmda, 9, 10, 11. 

horrida, 10. 

microstoma, 13. 

imirina, 12, 13. 

7iiuris variegati, 13. 

nana, 13. 

relicta, 10. 
Hyomoschus 

aquaticus, 460. 
Hyphantoniis 

hrachyptera, 477. 

capensis, 417. 

castaneo-fuscus, 477. 

nigriceps, 477. 

personata, 477. 

superciliosus, 477. 

i'OTi'or, 477. 

velatus, 477. 
Hypocbera 

nitens, 476. 
Hypocolius 

ampelimis, 473, 
HypopliEca 

chalyhea, 474. 
Hypotriorcbis 

chicquera, 504. 
Hypsipetes 

madellandi, 472. 
Hyrax 

capensis, 452. 

dorsalis, 452. 
Hystricbopsylla 

iripectinata, 391. 
Hystrix 

brachyura, 145. 

cristata, 144, 449. 

dorsata, 145. 

javanica, 449. 

longicaiida, 449. 

onacroura, 145. 

2)re}iensilis, 145. 

Ibis 

cBthiomciis, 510. 



Ibis 

berniert, 510. 

melanoccphalus, 510. 

strict ipeiinis, 510. 
Icterus 

aheillcei, 480. 

baltimore, 480. 

chrysocephalus, 480. 

jamaici, 480. 

spurius, 480. 
j tibialis, 480. 
j vulgaris, 480. 
I Icticyon 

venaticus, 440. 
j Ictonyx 
I frenata, 441. 
' lybica, 44] . 

z or ilia, 441. 
lonornis 
j martinic2cs, 525. 
! Isoniys 

variegatus, 448. 
I Itbaginis 
I geoffrnyi, 520. 
I Ixocincla 

I crassirostris, 53, 54, 60, 
I 82. 

lynx 

torqnilla, 487. 



Jaculus 

jac'uhis, 148. 
Jiilis, 215. 
Junco 

hyemalis, 480. 



Ketupa 

ceylonensis, 493. 

javanensis, 493. 
Kobus 

Z-o/^ 2. 

unctuosus, 2. 



Lacerfca 

(yf/i7zs, 22, 23, 24, 25, 
' 31, 32, 40. 
atlantica, 22, 26, 32. 
boettgeri, 25. 
cmrulea, 26. 
cam2}estris, 26. 
chlorogaster, 22, 25. 
defilippi, 25. 
derjugini, 22. 
(^^/^rey/i, 22, 24, 25, 28, 

32. 
echinata, 24, 25, SO, 

32. 



Lacerta 
fiunuma. 26. 
//w/fo/i, 22, 25, 26, 32. 
horvathi, 34. 
jacksoni, 19, 22, 23, 24, 

30, 32, 34, 35, 36. 
fey/.s, 22, 24, 26, 32. 
?///b/-(/«, 25, 31, 32. 
mosorensis, 24, 25, 26, 

27, 31, 34, 36. 
j*;Mr«fe, 22, 23, 24, 26, 

28, 31, 34, 37, 39, 
40. 

— bocagii, 36. 

— campestris, 25, 36. 

— melisselle'iisis, 26. 
■ — moiiticola, 36. 

— nigrivenlris, 22. 

— ser/j«, 22, 25, 26, 
36 

— tiliguerta., 25. 

— !'y;j?ra. 23, 25, 31. 
oceWfl/a, 22, 24, 26, 28, 

32. 
oxycej)liala, 24, 25, 26, 

27. 
peloponnesiaca, 2>1, 

40. 
pityusensis, 26. 
sardoa, 39. 
saxicola, 22, 25. 
schreiberi, 347. 
simonyi, 22, 24, 25, 26, 

32.' 
z'a«r2c-a, 37, 39, 40. 
vauereselli, 33, 34, 35, 

36. 
fiV/c^/s, 22, 24, 25, 26, 

32, 37, 39, 40. 

— gadovii, 347. 
i'm>am,23, 24,25, 31. 

Lagopus 

scoticus, 46, 100. 
Lagostoiiius 

trichodactylus, 450. 
Lagotbrix 

hmnboldti, 433. 
Lagotis 

cuvieri, 450. 
Lama 

glama, 150. 

huanacos, 460. 

paces, 150, 460. 

■peruana, 460. 
Lamprocoliiis 

aurat'us, 481. 

ehalyheus, 481 . 
Lampropsar 

aJjwes, 481. 
Lainprotornis 

(Biieus, 481. 



INDEX. 



Laniariiis 

quadricolor, 473. 
Lai'.ius 

collurio, 473. 

excuhitor, 473. 

lalitora, 473. 
Larus 

argeniatus, 77, 530. 

atricilla, 530. 

bricnneicephalus, 530. 

cachinnans, 530. 

caims, 531. 

cirrhocephalus, 531. 

dominicanus, 531. 

franMini, 531. 

fiisctis, 531. 

gelastes, 531. 

glaucus, 531. 

ickthyaetus, 531, 

leucopterus, 531. 

marimis, 531. 

novm-hollandice, 531. 

ridibundus, 77, 531. 
Latastia 

degeni, 30. 

hardeggeri, 30. 

neumanni, 30. 

pMllipsii, 30. 

spinalis, 30. 
Latax 

/MiJrzs, 138. 
Leggada, 169. 
Leipoa 

ocellata, 523. 
Lemmus 

lemmus, 146. 
Lemur 

alhifrons, 434. 

albimanus, 434. 

brunneus, 434. 

ca«a, 129, 434. 

coronatiis, 434. 

flavifrons, 434. 

fulvus nigrifrons, 5. 

— rufifrons. 5. 

macaco, 5, 434. 

mayottensis, 434. 

■mongoz, 434. 

ni^'er, 434. 

nigerrimus, 434. 

nigrifrons, 434. 

rufifrons, 434. 

rtifipes, 434. 

tarcligradus, 129. 

varius, 434. 

wfews, 129, 130. 

xanthomystax, 434. 
Leontopithecus 

midas, 128. 

cedipus, 127. 
Lepidosiren, 118, 119. 



Leptobradiium, 393. 

hasseUii, 393, 395, 4l.O, 
403. 
Leptoptila 

jamaicensis, 518. 

ochropiera, 518. 

rufaxilla, 518. 
Leptoptiliis 

argala, 509. 

crumenifcrus, 509. 

javanicus, 509. 
Lepus 

americanus, 451. 

brasiliensis, 146, 
451. 

capcnsis, 145. 

cunicuhos, 145. 

europcBUs, 451. 

nigricollis, 45 1 . 

sechioenensis, 180. 

sylvaticus, 451. 

timidus, 145. 

variabilis, 451. 
Leucodioptron 

canorum, 472. 
Leuconerpes 

candidus, 487. 
Leiicopternis 

pialliatiis, 502. 
Leuoosarcia 

picata, 518. 
Licmetis 

nasica, 495. 

pastinator, 495. 
Ligurinus 

chloris, 478. 

sinicus, 479. 
Limuocoi'ax 

niger, 525. 
Liinnotragus, 350, 353. 
Limosa 

agocephala, 529. 

lapponica, 529. 
Linota 

cannabina, 479. 

flavirostris, 479. 

rufescens, 479. 
Liothrix 

luteus, 471. 
Lobivanellus 

lobaius, 3, 530. 
Lomvia 

troile, 532. 
Lophsemus 

a?itarcticus, 516. 
Lophaetus 

occipitalis, 503. 
Lopbophorus 

impeyanus, 521. 

Ihuysi, 521. 

sclateri, 521. 



Lophufa 

ignita, 314. 
Loriculiis 

clirysonotus, 498. 

galgulus, 498. 

indicus, 498. 

pusilhis, 498. 
Loris 

gracilis, 434. 

volans, 129. 
Lovius 

domicella, 494. 
flavo-palliatus, 494. 

garrulus, 494. 

^ory, 494. 

tibialis, 494. 
Lota 

vulgaris, 229. 
Loxia 

bifasciata, 48(1. 

curvirostris, 480. 

Ic'ucoptera, 479. 

pityopsittacus, 479. 
Lucioperca, 215. 
Lutra 

felina, 181. 

leptonyx, 441. 

;«<?'ra, 138. 

jiazV, 441. 

vulgaris, 441. 
Ljcaoo 

pict'us, 440. 
Lycosa 

canescens, 300. 

Mabuia 

raddonii, 181. 
Maca^ciis, 121, 125, 126. 

arctoides, 431. 

assamensis, 431. 

brunneus, 4?1. 

cyclopsis, 431. 

cynomolgus, 431. 

ecaudatus, 126. 

i«?<a, 126. 

M?«<Ms, 126, 431. 

lasiotue, 431. 

leoninus, 431. 

maurus, 431. 

nemestrinus, 431. 

ocreatus, 431. 

pelops, 431. 

pJiilippinensis, 124, 129. 

pileatus, 431. 

rhesus, 431. 

rtifescens, 431. 

siieMMs, 126, 431. 

sinicus, 431. 

speciosus, 431. 

syricJda, 129. 

tcheliensis, 431. 



INDEX. 



XXXIX 



Machetes 

pugvax, 529. 
Machetornis 

rixoia, 484. 
Macropodus, 207, 215. 
Macropus 
agilis, 464. 
hennetli, 4()4. 
billanluri, 464. 
brachiuruii, 404. 
derbiaims, 4()4. 
(^orsa^/s, 464. 
giganteus, 464. 
melanops, 4(i4, 
parryi, 4(5 i. 
robustus, 464. 
rujicoUis, 464. 
rufus, 464. 
thetidis, 464. 
ualabatus, 464. 
Macropygia 

lepiogrammica, 517. 
2)ha^iancUa, 517. 
Man is 

dalmani, 124, lii3. 
pentadact(/la, 124, 

133. 
tricuspis, 463. 
(Pholidotis) qiganiea, 
463. 
Maimcodia 

chalybea, 482. 
Mareca 
penelojje, 514. 
sibilatrix, 514. 
Marmaronetta 

angustirostvia, 514. 
Marmosa 

dorsigera, 144. 
murina, 144. 
Marmota 

marmota, 147. 
ononax, 147. 
roteste, 169, 374, 375. 
Martes 

martes, 139. 
zibellina, 139. 
Megacepbalon 
maleo, 523. 
Megaderraa 

spasma, 131. 
Megalasma 

asiatica, 62, 490. 
hodgsoni, 490. 
virens, 490. 
Megalophrys 

(Leptobi'acbiuni) /(?(?, 

393-403. 
montaiia, 393, 396, 

399. 
Kaswte, 393, 396, 399. 



Melanerpes, 60, 76, 89. 
crythrucephalum, 487. 
supiercUiaru, 62. 
Melanocoryplia 
cakotdra, 484. 
mongolica, 484. 
gelfunieiisii', 484. 
Meleagvis 
gallopavo, 522. 
ocellata, 622. 
Meles 

ankama, 441. 
Icptorln/nchus, 441. 
meles, 140. 
taxus, 441. 
Melierax 

munogrammicus, 503. 
musicus, 5U3. 
polgzo?ius, 503. 
Mellivora 
capensis, 441. 
indica., 441. 
Melopelia 

leacoptera, 517. 
Melnphiis 

mdanictenis, 480. 
Melopsittacus 

und'iclatus, 45, 85, 
499. 
Melursus 

ursinus, 442. 
Memira 

su-perha, 484. 
Mephitis 

mephitica, 181, 441. 
Mergus 

albellus, 514. 
merganser, 514. 
serrator, 514. 
Meriones, 6. 
Merops 

apiaster, 4S9. 
Mesoplodoa 
australis, 422. 
bidens, 424. 
densirostris, 421, 422. 
haasti, 422. 
(Dioplodon) <p'ayi, 
420. 421, 422. 
Mesopsylla 
exwta, 12. 
Meta 

argentata, 298. 
Metachirus 

opossum, 143. 
Metopiana 

peposaca, 514. 
Metriopelia 

melano-ptera, 518. 
Microcebus 
smithi, 434. 



Micrnglossus 

atcrritiius, 494. 
Microsittace 

ferriiginea, 496. 
Minrotus 

arvcdis, 173, 175. 

calamorum, 174, 175. 

MC^", 176. 

irene, 173. 

limnophilus, 174, 175. 

malcolmi, 173, 174, 175, 
178. 

?;M.r, 176. 

ratticeps, 175. 

sai'iY, 170. 

(Anteliomys) cliinensis, 
175. 

(Caryomys) eya. 175. 

(Eothenomys) melano- 
gasier, 176. 
Midas 

ckrysomelas, 433. 

geoffroii, 433. 

labiatus, 433. 

7)iystax, 433. 

cedipus, 433. 

rosalia, 433. 

rufimanus, 433. 

ursulus, 433. 
Milvago 

australis, 504. 

chlmachima, 4, 504. 

chimaiigo, 504. 

megalopterus, 504. 
Milvulus 

tyrannus, 484. 
Milvus 

cBgyptius, 504. 

govinda, 504. 

iciinus, 504. 

migrans, 504. 
Mimus 

polyglottus, 471. 

saturinus, 471. 
Mirafra 

affinis, 484. 

caniillans, 484. 
Miro 

albifrons, All. 
Mirounga 

leonina, 133. 
Mitua 

tomentosa, 523. 

tuberosa, 523. 
Molotbrus 

badius, 481. 

bo7iariensis, 481. 

purpurascens, 481. 
Moniotus 

brasiliensis, 489. 

subrufescens, 489. 



xl 



IXDEX. 



Monachus 

albiveiiter, 44'>. 
Monodoii 

monoceros, 155. 
Monticola 

ct/aMus, 470. 

saxatilis, 470. 
MoutitVingilla 

nivalis, 479. 
Morphnus 

guianensis, 502. 
Moschus 

moschi ferns, 150, 
458. 

sifariicus, 180. 
Motacilla 
flava, 472. 

hcguhris, 472. 

■melanope, 472. 

raii, 472. 
Motella, 215. 

mediterranea, 207. 
Mungos 

ichneumon, 136. 
Mania 

castaneithorox, 101. 

ferruginea, 475. 
flaviprgniJia, 101. 

■maja, 476. 

malabarica, 476. 

malacca, 476. 

nisoria, 476. 

■punctularia, 476. 

ruhro-nigra, 476. 

striata, 476. 

topela, 476. 
Muraina, 216, 219. 
Mus 

amphihius, 147. 

arboricola, 448. 

avellanarius, 147. 

bandicota, 448. 

harharus, 448. 

blanfordi, 448. 

cervicolor, 448. 

cricetus, 147. 

exula7is, 448. 

gansuensis, 169. 

jaculus, 148. 

Icmmus, 146. 

lepori^ms, 146. 

ioiigipes, 148. 

marmota, 147. 

minutus, 448. 

monax, 147. 

musculus, 122, 146; 
147. 

porcelhis, 146. 

rff^MS, 4, 147, 448. 

tftriatus, 148. 

sylvaticus, 148, 448. 



Mus 

ferresirh. 147. 

I'otois, 148, 149. 
Muscardimis 

avellanarius, 147, 447. 
Muscicapa 

atricajnlla, 473. 
Mii«ophaga 

rosscs, 3. 

violacea, 491. 
Miistftla 

harbai-a, 138. 

hedfordi, 169. 

dorsalis, 169. 

erminea., 138, 139, 
441. 

fl.avigula, 181. 

foina, 441. 

y^ro, 139. 

(7?/fo, 138. 

'kathiah, 168, 169. 

i?««2?m, 138. 

luiris, 138. 

mariJes, 139, 441. 

nivalis, 168. 

j-yennanti, 441. 

putorius, 139, 441. 

ri.TOda, 168, 169. 

russelliana, 168. 

vulgaris, 138, 441. 

zibellina, 139. 
Mustelis 

diver sis, 122. 
Mycerobas 

■melanoxanthus, 478. 
Mycetes, 433. 
Mycteria 

americana, 509. 
Mviopliones 

'horsfwldi, 470. 
Mynogleaes 

c/iz'to«j, 299, 308. 
309. 

insolens, 297. 

■marrinen, 310. 
Myodes 

leinmus, 448. 
Myopotamus 

coypu, 449. 
Myopsittacus 

ononachtis, 4-96. 
Myospalax 

ca>isi(s, 178, 179. 

— shenseius, 178. 

fontanieri, 391. 
Myotalpa 

rufescens, 179. 
My Otis 

altarium, 161. 

bechsteini, 161. 

vwupinensis, 162. 



j Myotis 

j pequinius, IGl. 

' Myoxus 

dry as. 447. 

(?//s, 447. 
I quercinus, 447. 
i Myristicivora 
I luctuosa, 516, 
' Myrmecopiiaga 

didacfyla, 132. 

yw/jato, 124, 132, 402. 
; ietradae.t.yla, 133. 
{ tr'idactyla, 124, 132. 

Myrus, '113. 189, 192, 
' 193, 216, 217, 218. 

219. 

vulgaris, 191, 192. 
Myzantha 

garrula, 474. 



Naia 

tripudians. 4. 
Nandinia 

binofata, 439. 
Nanodes . 

discolor, 499. 
Nasua, 137. 

narica, 442. 

nc/a, 442. 
Necator, 549. 
Nectnrus, 103, 105, 115, 

119. 
Nemorboedus 

goral, 456. 
Keopbema 

6o(«-/.7V, 499. 

chrysogastra, 499. 

elegans, 499. 

pulchella, 499. 

splendida, 499. 

venusta, 499. 
Neopbron 

percnopterus, 504. 
Neopsylla 

ffiicwff, 388. 390. 

hid.entatiformis, 387, 
.388. 

co7npar, 387. 
Neotetracus, 159. 

sinensis, 162. 
Neotragus 

pygmea, 152. 
Nephila 

argentatum, 298. 

venosa, 299. 
Neropbis, 183, 201, 219, 
225, 235. 

(equo7it(s, 199, 200, 
248. 



xli 



NesocVien 

samhiveiids, 5 1 'J. 
Nesonetta 

cmcJdttndica, 514. 
Nesopelia 

galapagoensis, 517. 
Nestor 

noiabilis, 84, 491. 
Ninox 

hoohook, 71, 72, 493. 

novce-scelandia, 493. 
Nippouia 

temmincJci, 510. 
Nisaetiis 

fasciafus, 502. 

pennatns, 5t)2. 

f^pi log aster, 502. 
Noetilio 

leporinus, 131. 
Nothocrax 

urunmtum, 523. 
No til lira 
. maculosa, 533. 
Notiosorex, l(i7. 
Niioit'raga 

caryocatades, 482. 

NUCTUS 

delalandii, 30. 
Numenius 

arquata, 529. 

femoralis, 529. 

phcBopus, 529. 
Nurairla 

^//i'oifi, 522. 

meleagris, 522. 

mitrata, b'2'2. 

pfilorhgncha., 522. 

reiehenoiui, 522. 
Nyctea 

scandiaca, 493. 
Nycticebus 

javanicus, 434. 

tardigradics, 434. 
Nycticorax 

caledonicits, 508. 

gardeni, 508. 

griseiis, 508. 

violaceus, 4, 508. 
Nyctidromus, 60. 
Njctipithecus 

feliims, 433. 

rufipes, 433. 

trivirgatus, 433. 

vociferans, 4.33. 
Nyinpliicus 

cornufus, 499. 

uvceensis, 499. 



Ocbotona 
cansa, 180. 

Proc. Zool. Suc- 



-191 



Ocliotona 

hodgsoni, ISO. 

tihelana, 180. 
Ocjtodoii 

cumingi, 449. 
Ooycli'oimis 

australis, 525. 

e(/r/«, 525. 

Juifcus, 52.5. 

I'lfresiia.f/anns, 525. 

sglvestris, 525. 
Ocypliaps 

iophoies, 518. 
Odobenus 

rosmancs, 134. 
Odontopborus 

dentatus, 521. 

guianensis, 521. 
CEdeniia 

nigra., 514. 
QCdicnennis 

hlstriatits, .529. 

grallarms, 77. 529. 

scolopax. 76, 77, 529. 

nuperciliaris, 529. 
Q£na 

capoisis, 518. 
Onyehogale 

frena/a, 464. 

unguifera, 464. 
Opbichthvs, 183, 189, 
219. 

(Rpbasgebrancbus) nw- 
6e»V«.?, 192, 193, 216, 
217, 245. 

(Opbisuriis) serpens, 
193, 216. 
Ophidium, 183, 205, 
206, 214, 215, 216, 
217. 219, 220. 

harhafum. 197, 204. 
210,247. 
Orcbesticiis 

fl^'er, 47.5. 
O rein us 

orca, 1.58. 
Oreas 

ca.nna, 456. 
Oreofrngiis 

snltator, 455. 
Oriolns 

indicits, 473. 

hmdoo, 473. 

melanocep>halus, 473. 
Ornithorhyuebiis 

paradoxus, 181. 
Ortalis, 58. 

alhiventris, 523- 

ga,mda, 523. 

motmot, 523. 

ruficauda, 56. 523. 

1, No. XXXVIII. 



Oftali.'i 

vetida, 523. 
Ortyx 

virginiaiius, 521. 
Orycteropiis 

(Bthiopknis, 463. 

capensis, 4(]3. 
Oryctolagiis 

cunicidu.s, 145. 
Oryx 

algazel, 1. 

heatrix, 456. 

fieisa. 456. 

gasella,, 152. 

leucoryx, 4.56. 
Oryzoboriis 

crassiroslris, 478. 

torridus, 478. 
Osiuotrerori 

hicincia, 516. 

.vernans, 516. 
Ostinops 

decumanus, 480. 
Otaria 

calij'orniana, 443. 

hookeri, 443. 

juhata, 443. 

pusilla, 443. 
Otis 85. 

tarda, 73, 526. 
Olocorya 

alpestrls, 4S4. 

hU.opha, 484. 
Otocyun 

megalotis, 440. 
Ouacaria, 433. 
Ourebia 

montana. 1. 

nigricaudata, 2, 455. 
Ovis 

a.mmon, 153. 

aWgs, 153. 

hlanfordi, 457. 

hurrhel, 457. 

cyeloceros, 457. 

giiineensis, 153. 

hodgsoni, 457. 

musimon, 457. 

strepsiceros, 15-3. 

tragelaphus, 457. 

vignei, 457. 



Paoliynns 

hruchyurus, 2, 497. 
Padda 

oryzivora, 476. 
Pagellus, 215. 
Pagopbila 

eburnea, 531. 
Palajopsylla, 388. 

38 



xlii 



INDEX. 



Palieornis 

alexandri. 498. 
caniceps, 498. 
cyanocephala, 498. 
docilis, 498. 
fasciafa, 498. 
lovgicmida, 498. 
nicoburioa, 498. 
peristerodes, 498. 
rosa, 498. 
torquatus, 498. 
Palaiiiedea, 86. 
cor nut a, 611. 
Pandion 

haUaetus, 501. 
Piimirus 

hiarmicus, 471. 
Papio 

cynmnolgos, 128. 
hamadryas, 127, 
128. 
Paradisea 
minor, 482. 
raqgiana, 60. 
ri<6m, 2, 482. 
Paradisorois 

rudolphi, 60. 
Paradoxurus 
aureus, 438. 
grayi, 438. 
hermaphroditus, 438. 
larvatus, 438. 
leuGomystax, 438. 
52(^t3r, 438. 
jjalla&i, 438. 
phi/ippensis, 438. 
preheiisilis, 438. 
Pardosa, 298. 

canescens, 300. 
Pareiasaiinis, 118. 
Pareudriliis, 413. 

paUidus, 419. 
Paroaria 

capitata, 477- 
ciocullata, 477, 485. 
larvata, 477. 
Parus 
ai'er. 471. 
boreaiis. 471. 
C(srulaus, 471. 
caudatus, 471. 
cridatus, 471. 
cyanus, 471. 
major, 471- 
paluntris, 471. 
varius, 471. 
Passer 

aUirio, 478. 
arcuatus, 478. 
diffumis, 478. 



Passer 

biteus, 478. 
montanus, 478. 
salicicola, 478. 
simpilex, 478. 
Pastor 

roseus, 482. 
Paiixis 

galeata, 523. j 

Pavo , 

cristatus, 522. 
nigripe/mis, 522. 
spicifer, 622. 
Pelecaiius, 83. 

conspic'illatus, 506. 
cribpus, 506. I 

fuscus, 506. 

manillensis, 506. ' 

mitratus, 50('). 
onocrotahis, 606. 
rufescens, 501 >. 
trachyrhynchvs, 3. 
Pelobates, 394. 
Pelodjtes 

pimdatus, 397. 
Penelope i 

cristata, 523. j 

greeyi, 523. i 

jacucaca, 623. | 

marail, 623. ' 

pileata, 523. ! 

purjmrascens, 623. 
supercUiaris, 58, 523. 
Penthetria 
I laticauda, 4. 
I Peragale 

lagot/s, 466. 
Peraiueles 

ohesula, 466. 
Perca, 183, 210, 215, 
216, 219, 227, 237, 
240. 
fluviatilis, 214, 229, 
248. 
Perdicula 

arguundah, 621. 
asiatica, 521. 
Perdix 

cinerca, 520. 
Jiodgsoni(s, 520. 
Periobasta, 412. 
Peristedion 

catapliracium, 201, 202, 
210, 246. 
Peris tera 

geoffroyi, 518. 
Peristetlius, 183, 195, 
201, 203, 216, 219. 
cataphradus, 201, 202, 
210, 219, 246. 



Pornis 

apiooruH, 601. 
ptilorhyncha, 501. 
Perodicticns 
/jo«o, 4.34. 
P eta urns 

australis, 464. 
bret'iceps. 464. 
sciureus, 464. 
Pehrogale 

brachyoiis, 464. 
peniciUaia, 464. 
xanthoinis, 464. 
Petronia 
pdronclla, 478. 
si;Mftfl, 478. 
Phacoelioenis 
(sthiopiouE, 461. 
africaniis, 461. 
Pbaettiusa 

■magnirostrls, 530. 
Phalacrooorax, 83. 
a.fricanun, 507- 
brasillanus, 507. 
c«rfto, 507. 
carimculatus, 507. 
gracului, 507. 
javnuicus, 507. 
higubris, 507. , 
pygincB'us, 3. 
varius, 507. 
Pbaps 

chalcoptera, 618. 
elcgans, 618. 
hidrionica, 518. 
Pliascolarctos 

cinereus, 464. 
Pbascolomys 
latifrons, 465. 
rnitchclli, 466. 
Pbasianus 

colchicus, 621. 
decoUatus, 521. 
ellioti, 621. 
jjrincipalis, 521. 
reevesi, 621. 
scemmerringi, 521. 
torqticdus, 521. 
versicolor, 521. 
•wcdlicMi, 521. 
Pheretima, 412, 418, 
419. 
cingulata, 413. 
';«o?;itt«a, 413, 414, 416, 
417. 
Pbeucticus 

chrysogaster, 478. 
Pblogcenas 
lueonica, 618. 
s/airi, 618. 



INDEX. 



xliii 



Phooa 

Iconina, 133. 

rosmarus, 134. 

ursina, 133. 

vifuliua, 133, 134, 
443. 
Phoetena 

■jihoccena, 158. 
Phodilus 

badius, 492. 
Plioenicoptenis 

ignipaUiatus, 511. 

roseus, oil. 

rubra, 511. 
Pbonipara 

canora, 477. 

olivacea, 477. 
Phoronis, 233. 
Phrygilus, 477. 

alaudimis, 478. 

iruticeti, 478. 

f/a^f, 478. 
Plijseter 

catodon, 157. 

macrocephal'us, 157. 

microceplialus, 157. 

microps, 157. 

tursio, 157. 
Pica 

hootanensis, 482. 

mauritanica, 482. 

rustica, 482. 

sericea, 482. 
Pinicola 

enucleator, 479. 
Pionopsittacus 

pileafus, 497. 
Pi onus 

ohalcopterus, 497. 

pisciis, 497. 

maximiliani, 497. 

menstruus, 497. 

senilis, 497. 

sordidus, 497. 
Pi pile 

cujtihi, 523. 

cumanensis, 523. 

jacutinga, 523. 
Pipilo 

erythroiMlia Imu s, 4 80. 
Pilangus 

sulphuratus, 484. 
Piihecia, 433. 
Pitbecus, 121, 125. 
Pitta 

hengalcnsis, 484. 

strepifans, 484. 
Pitvmys. 173. 
Pjatali-a 

r///jr/, 510. 

h ucorodia, 510. 



Platycerciis 

ad da idee, 498. 

hoA-nardi, 499. 

browni, 499. 

elegans, 499. 

eryihropepliis, 499. 

eximius, 499. 

jlaveolHS, 84, 499. 

flavivc72iris, 499. 

icierotls, 499. 

pallidiceps, 499. 

pi.leatus, 499. 

semitorquafus, 499. 

S07iarius, 499. 
Plecotus 

(Egyptiacus, 160. 

«r/e/, 160. 

awnYew, 131, 160, 445. 

clivistiei, 160. 

homochrous, 160. 

sac7-imontis, 160. 

Wflr^^/, 160, 161. 
Plectrophenax 

nivalis, 480. 
Plectropterus 

gambensis, 512. 

niger, 512. 

rueppelli, 512. 
Plegadis 

falcinellus, 510. 

gnarauna, 510. 
Pl'>ceus 

atrigula, 477. 

bengtdensis, 477. 

wanyar, 417. 
Plotns 

ank/7iga, 3, 507. 

leraillanti, 507. 

melano g aster , 507. 
Pluvianus, 73, 76, 77, 
89. 

cegypiius, 74, 75. 
Podager 

nactinda, 314. 
Podargus 

cMyfm, 58, 60, 489. 
Podiceps 

cristaius, 78, 81, 
532. 

griseigena, 532. 
Pcecilonetta 

bahamcnsis, 514. 

erythrurhyncha, 514. 
Poeoceplialiis 

fusciaqjillus, 497. 

fuscicollis, 497. 

gulielmi, 497. 

meyeri, 497- 

robust us. 497. 

raepprlli. 497. 

smrqnlti^, 497. 



Poepbngiis 

grminiens, 454. 
Poepbila 

aciUicauda, 476. 

cincia, 476. 

gouldice, 476. 

mirahilis, 476. 
Polioaetiis 

plitmbetts, 502. 
Poliopsar 

andamanensis, 481. 

ftZ^;'/</, 481. 

malaharicus, 481. 
Polyacantbus, 207, 

215. 
Polyboroides 

typicus, 504. 
Puly boras 

hrasiliensis, 504. 
Polyplectron 

bicalcaraf.nm, 522. 

chinquis, 522. 

germaini, 522. 
Polytelis 

alexandrcB, 498. 

barrabaudi, 498. 

melanurus, 498. 
Porcula 

salvania, 461. 
Poromera 

./brcM, 30. 
Porpbyrio 

a/iws, 82. 

caruleus, 525. 

calvus, 525. 

edmardsi, 525. 

madagascarieiisis, 



82, 



melanonofus, 82, ,525. 

poliocephalas, 525. 

vitiensis, 525. 
Porrbothe.le 

antlpodiana, 298. 
Porzaiia 

Carolina, 525. 

maruetta, 525. 

notata, 525. 
PotamoL'boeriis 

africa.mis, 461. 

penicillatus, 461. 

porcHs, 140. 
Potorous 

tridactylus, 464. 
Prntiiieola 

ruhetra, 471. 
Presbytis, 124. 

aygula, 124, 127. 

mitratus, 124, 127. 
Procyon 

cancrivorus, 442. 

fo/r-r, 140, 442. 



xliv 



ProRflromys, 159, 174, 
177. 
hedfordi, 159, 177. 
Proslheiiiadora 

novcB-ze.alandia, 474. 
Proteles 

cridafus. 4-'i9. 
PTOt.opteriis, 118, 119. 
Psainmouiys 

ohesus, 448. 
Psephotiis 

chri/sop''er];gms, 499. 

hcematonotus, 84, 
499. 

multicolor, 499. 

'pulclierrimiis, 499. 
Pseutlis, 396. 
Pseiidochinis 

•perccjrinus, 464. 
Pseudoleistcs 

virescens, 481. 
Pseiulotantalus 

?7«-s. 5(19. 

leucocephalus, 509. 
Psiltacula 

passemia, 496. 
P.sittaciis 

eniJiaciis, 497. 

fivnic/i. 497. 
Psittenteles 

chloroleindotus, 494. 
Psittinus 

incerius, 498. 
Psophia 

crepitcnis, 527. 

leacoptera, 527. 

viridis, 527. 
Pternisle.^ 

a/pr, 520. 

irifiiscaius. 520. 

leucoscepus, 520. 

nudicol/is, 520. 

gwainsoni, 520. 
Ptei'oules 

alchata, 519. 

arenarms, 519. 

hicinctvs, 519. 

ea'UHfus. 519. 

fasciatus, 519. 

namaqiia, 519. 

sener/aJh/.% 519. 
PteroglossLis 

ivi^eripf'HS, 490. 

lorquaius, 490. 

vnedi. 490. 
Pteroniys 

alho-nifus, 446. 

inornatus, 446. 

Jeiicogenys, 446. 

magiiificus, 446. 

orrt/, 446. 



Pteropus 

er/w/is, 180. 
fo7-mosus, 444, 
gouldi, 444. 

medius, 444. 
2)oliocephalns, 444. 
pse/apkon, 444. 

scapulatus, 444. 

varivpi/rus, 130. 
Ptilonorhynehus 

smiihi, 482. 

violaceus, 60, 482. 
Ptilopnchys 

centralis, 520. 
Ptiiopus 

hdl'iis, 85. 
fasckdas. .516. 
jamhu, 516. 

melavoceplialus 516. 

porphi/raceics, 516. 
Ptilorliis 

paradisea, 482. 
Ptilostomns 

scnegaleiisis, 483. 
Ptistes 

eryfhropterus, 493. 
Pucrasia 

darwini, 521. 

macrolopha, 521. 

A'«?i !*/; oapila, 52 1 . 
Pudua 

himilis, 459. 
Puflinus 

avgJorum, 531. 

Jciihh, 531. 

major, 531. 
Piilex 

iri'itans, 5. 
Pulsatrix 

torquaia, 493. 
Pjcnonotus 

atricapHlus, 472. 

crocorrhous, 472. 

luemorrlious, 472. 

jocosus, 472. 

Icucotis, 472. 

sinensis, 472. 

xmithopyqos, 472. 
Pyf;atlirix,'l24, 127, 128. 
Pygosceles 

tmnkdus, 532. 
Pyi-aiiga 

sairu, 475. 
Pyrenestes 

albifrons, 477. 
Pyroiiielana 

aurca, 476. 

franciscana. All. 
Pyrrbocoiax 

cdpivv!', 483. 

gracuhi.^. 483, 



Pyrrhula 

eiiropaa, 479. 

ruhicilla, 479. 
Pyrrhiilauda 

verticals, 484. 
Pyrrhulopsis 

personata, 498. 

splfndens, 498. 

tahvensis, 498. 
Pyrrliiii-a 

crueniafa, 496. 

leucotis, 496. 

perlata, 496. 

OTY/ff7'«, 496. 
Physeter 

catodon, 124. 

macrocephalus, 124. 
Pytelia 

citerior, 475. 

phoenicoptera, 475. 

wieneri, 475. 



Quelea 

occidentalis, 477. 

»'?<ssf, 477. 

sanguinirosh'is, 477 . 
Querqiiedula 

braHiliensis, 514. 

castanea, .514. 

circia, 514. 

crecca, 514. 

ci/anoptera, 514. 

faJcata, 514. 

flavirostris, 514. 

fm-musa, 514. 

gihherifrons, 514. 
Qiiiscalus 

luguhris, 481. 

versicolor, 481. 



Rallina 

pcecilopitera, 625. 
Eallua 

cdihoti, 82. 

aqiiaticus. 525. 

celeherisis, b'2b. 

mandatus, 5'lb. 

pectorcdis, 525. 

rhytirhynchus, 525. 
Rana. 40i. 
Rangit'er 

tarandn^, 151, 459. 

— caribou, 181, 182. 
Raphicenis 

onelavoiis, 455. 

tragnluR, 455. 
Recnryirostra, 65. 

fivocefla. 77, 78, 529, 



IXDEX. 



xlv 



Rhamphastos 
ariel, 63, 489. 
brevicarinatus, 489. 
carlnalus, 63, 489. 
cuvieri, 489. 
discolonis, 489. 
erythrorhynchus, 490. 
!;oca,r<?, 490. 
("(W, 490. 
viiellinus, 490. 
Eliampbocoelns 
hrasilius, 475. 
Pliaiiiphocorys 
clotheyi, 484. 
Ehea, 90, 9.3. 

americana, 51, 52, 

54, 535. 
dariuiiii, 535. 
■inacrorhyncha, 535. 
Ehinoceros 

bicornis, 144, 

453. 
lasiotis, 453. 
soudaicus, 453. 
sumatroisis, 453. 
unicornis, 144, 
453. 
Ehinochetiis 

jubatus, 528. 
Rliinodenna 

darivini, 401. 
Ehiiioloplius 

Cfr-H ?r/7<s ^j'i^ ;?? ///^,.s, 
11)0. 
Eiliizoinys 
badius, 449. 
(^avfA, 179, 180. 
sinensis, 179. 
vestitus, 179. 
EliodoTiessa 

caryophyllacea, 512. 
Eliynchotus 
perdicarius, 533. 
rufesceiis, 533. 
Rhylicloceres 
plicatus, 488. 
undulaius, 488. 
Eissa 

tridactyJa, 531. 
liollulvis 

cristatus, 521. 
Ro.'trhamus 

sociaiiilis, 504. 
Rupicapra 

rwpicapra, 152. 
tragus, 466. 
Eupicola 

crocea, 2, 484. 
Ruticilla 

phoenicHfii.s, 471. 
CiM//.s, 471. 



Saiinii'i 
mor^a, 128. 
sciurea, 129. 
Salaniandra 
rtifra, 347. 

maculosa, 823, 324, 
325, 326, 328, ^29, 
330, 33], 332, 333, 
340, 342, 347. 

— a/^/ra, 324, 330. 

— coccinea, 338. 

— cors/m, 324, 330, 
340, 342, 343. 

— europcsa, 324. 

— qallaica, 327, 333, 
334, 335, 33!i. 

— y«o//c/-i, 324, 327, 
333, 334, 335, 336, 

337, 342, 347. 

— nigriveniris, "324. 

— quadrivirgnta, 32i. 

■ — tcsniata, 324, 327, 
330, 333, 336, 337, 

338, 339, 345, 346, 
347. 

Saltator 

aurcmtiifostris, 475. 

magnus, Alb, 

simllis, 475. 
Sarcidiornis 

cnrtmrulata, 512. 

mdanoH.ok(,, 3, 
512, 
Sarciophorus, 89. 

pectoralis, 76, 529. 
Sarcophilus 

satanicus, 100. 

ursinus, 466. 
Sarcorbaiupluis 

cequatoricdis, bQf). 

gryphus, 2, 505. 
Sargiis, 183, 208, 215, 
219. 

rondeletii, 209, 210, 
211, 216, 248. 
Sauropatis, 64. 
Saxicola 

ananthe, 471. 
Seal ops 

aqtudicus, 143. 
Sc.ipfceir.i 

Jcnoxii, 31. 
Scardaf'ella 

squamosa, 517. 
Scauriis 

striat'us, 10. 
Sceloglanx 

albifacies, 493. 
Scitena, 215. 
Soiui'opterus 
Jimhriatus, 446. 



Sciuropterus 

layardi, 446. 

russicus, 149 

volans, 148, 446. 

volucelia, 446. 
Sciurotainias 

davidianus, 169. 

— cousobrinus, 169, 
371, 38i, 385, 
393. 
Sciurus 

<estivans, 44.5. 

arisoiiensis, 44.5. 

atrodorsalis, 445. 

bicolor, 445. 

caniceps, 445. 

castaneiveiitris, 445. 

ciiiereus, 149, 445. 

dorsalis, 445. 

ephippmm, 445. 

ergihropus, 445. 

flaviis, 149. 

getidus, 149. 

griseo-Jlavus, 445. 

gri-<eiis, 445. 

hudsonicus, 445. 

hgpopyrrkus, 445. 

iudicus, 445. 

ludovicianus, 445. 

macrurus, 445. 

madri.gascaricnsis, 445. 

mixijnus, 445. 

niger, 149. 

nigro-vittatus, 445. 

palmarum, 445. 

pla)itani, 445. 

Ijrevasti, 4i5. 

punctatus, 2. 

pusdlvi. 445. 

pygerythrus, 445. 

sframineus, 445. 

striatus, 149, 

syrictciis. 445. 

fennenti, 445. 

tristriatus, 445. 

variabilis, 446. 

variegafus, 446. 

volans, 149. 

vulgaris, 118, 446. 

vulpina, 446. 
Scolopax 

rusticola, 529. 
Scops 

asio, 493. 
^««, 493. 
glabripes, 493. 

lempiji, 493. 

leucotis, 493. 
senegalensis, 493. 
Scopus 

nmhrctfa, 3, .509. 



xlvi 



INDEX. 



Scoto])elia 

bouvieri. 71. 

pell, 493. 
Scytbrops 

nov(B-hoUandi<p, 66. 
Selenidera 

onamdirostris, 490. 
Seleucides 

niyrirans, 482. 
Seoinopithecus 

ccphalopterus, 430. 

entellus, 43a. 

fasdatus, 430. 

hi/poleuciis, 430. 

mmirus, 430. 

ohscuriis, 430. 
Sericiilus 

melinus, 473. 
Serinus 

aiigolensis, 479. 

canariiis, 479. 

canicollls, 479. 

horhdanus, 479. 

i'o;'i;?<s, 479. 
Serpentarius 

rcjytilivorus, 505. 
Serrainis, 183, 215, 219. 

cabriUa, 2U9. 
Sialia 

wilsoni, 471. 
Sibia 

cap i drat a, 472. 
Siiiiia 

(Bihiops, 128. 

apcdia, 126. 

cpe/fe, 128. 

aygula, 127. 

capHcina, 128. 

cephus, 127. 

ct/namolgus, 128. 

f^Mwa, 127. 

faunus, 12'). 

hamadryas, 127. 

jacclms. 127. 

mid.as, 128. 

morta, 128. 

nemrea, 127. 

aedipns, 127. 

paniscus, 127. 

«(/'?/r«s, 121, 122, 124, 
126. 429. 

sriurea, 129. 

silenus, 126. 

fiphitf.v, 126. 

s^mwa, 121, 122, 124, 
125. 126. 

ayriolita, 129. 
Siphonostoma, 183, 219. 

rnndelciii. 200, 246. 

Av/;///^. 200, 201. 246. 
Sireilon, 103. 



Sitagra 
' ^«<ec//rt, 477. 
Sitta 

c(Bsia, 472. 
Smaris, 183. 

»«««/•«, 203, 207, 208, 
217, 219, 247. 

vulgaris, 203. 207, 208, 
216, 217, 219, 247. 
Smilodon 

calif ornic us, 181. 
Solaster 

endcca, 45. 
Somateria 

moliissima, 514. 
Sorex, 167. 

aquaiicus, 143. 

araneus. 143, 164. 

bedfordice, 164, 
16 X 

cristaius, 143. 

cylindricauda, 164. 

minutus, 164. 

guadraticmida, 167. 

xvardi, 165. 
Sori cuius 

caudutus, 165, 
166. 

onacrurus, 165, 166. 

sacratKs, 165. 
Spathopteriis 

alexandrcB, 2. 
Spatula 

clypenta, 514. 
Speutyto 

cunicularia, 493. 
Spennest es 

cucuUata, 475. 

fringilloides, 475. 

nana, 457. 
Spermophila 

albogidaris, 478. 

ccBridescens, 478. 

collaria, 478. 

culeri, 478. 

guttvralis, 478. 

hypoleuca, 478. 

lineata, 478. 

lineola, 478. 

nigro-aurantia, 478. 

plumhea, 478. 

torqucola, 478. 
Spermophilus 

r7Y?//«,';, 446. 

grammarus, 446. 

mcxicanus, 446. 

oriongolicus, 446. 

fridccimlineaivs. 446. 
Sperniospiza 

gi(i/rit((, 477. 

hcemnthna, 477. 



Spliagolobus 

atratus, 488. 
Spheniseus 

demersus, 532. 

hundwldti, 532. 

magellanicus, 632. 
Spbenocercus 

sphenttrns, 616. 
Sphingurus, 449. 

insidiosiis, 450. 

mexicamis, 450. 

prehi'.nsilis, 450. 

spinosus, 4.50. 

villosus, 460. 
Spilogale 

pidorius, 137, 181 . 
Spiloi'iiis 

fecZ/ft, 503. 

c/;eefe, 503. 

spilogaster, 603. 
Spiroptera 

Uganda, 554, 555. 
Spiziietus 

hellico&us, 68, 69, 
502. 

caligatus, 503. 

ccyloiiensis, 503. 

coronalus, 2, 503. 
■ nipalensis, 503. 

orientalis, 503. 

ornat'us, 503. 

tyrannus, 503. 
Spizella 

socioJis, 480. 
SquaLarola 

helvetica, 529. 
Starnoenas 

cyanocephala, 518. 
Steatoniis 

caripensis, 489. 
Stenoponia, gen. nov., 
391. 

cc8/esif/s, 391, 392. 

tripeclinaia, 391, 392, 
393. 
Stephanophorus 

leucocephalus, 474. 
Stercurarius 

antarcticus, 630. 

biiffoni, .530. 

crepidaius, 630. 

7J077? atorliinus, 530 . 
Sterna 

cantiaca, 530. 

fluviatilis, 78, 630. 

liirundo, 78. 

minuta, 530. 
Strepera 

anapkonctifisi, 483. 

J'uliginosa. 483. 

grnndina. 483. 



Strepsieeros, 349. 
huxfoni, 348. 
capensis, 348. 
imherbix, 456. 
kudu, 456. 
Strepsilas 

inferpres, 529. 
Stringops 

hahi'optilus, 83, 85, 
499. 
Strix 
jlammea, 71, 72, 

492. 
ferlata, 71, 72. 
punctatlssima, 1'2. 
Strongylus 

equinus, 553. 
Struthidea 

cinerea, 473. 
Struthio, 51, 59, 
93. 
camehis, 535. 
mol)/hdopJiaiies, 55, 
535. 
Sturnella 
defilippi, 481. 
ludovicicma, 481. 
SLurnopastor 
contra, 481. 
Sturniis 

meiizhieri, 481. 
vulgaris, 481. 
Sula 

bassana, 507. 
letccogastra, 507. 
piscator, 507. 
serrator, 607. 
Suricata 

teiradactyla. 439. 
Siirnia 

funerea, 493. 
Sus 

andamanensis, 461. 
bahi/russa, 140. 
cristatus, 461. 
guineensis, 140. 
leucoviystax, 461. 
papuensis, 461. 
parens, 140. 
acrnfa, 140, 461. 
to jam, 140. 
Svcalis 

arvensis, 479. 
flaveola, 60, 479. 
luteola, 479. 
Sylvia 

atricapUla, 471. 
cinerea, 471. 
curriK-a, 47 1 . 
sylmcola, 471. 
truchiluK, 471. 



IXJJKX. 




xl 


Sylvilagiis 




' Tanagra 


hrasUiensis, 146. 




striata, 474. 


Syraphalangus 




Taaagrella 


synlacfi/lus, 4, 


353, 


cyunomclcBna, 474. 


356, 357, 358. 




•«;<'//«, 474. 


Syngnathiis, 183, 


194, 


Tantalus, 322. 


195. 196, 200, 


202, 


looulator, .509. 


203, 204, 201), 


212, 


Taaygnatliua 


215, 216, 217, 


219. 


lu'onensis, 498. 


acMS, 193, 218, 


219, 


megalorhynchus, 498 


245. 




muelleri, 498. 


Synoecus 




Tapir us 


austral is, 521. 




americamis, 453. 


Syrnium 




indicus, 453. 


«/mco, 71, 492. 




terrestris, 155. 


indranee, 492. 




Tatu, 124, 141, 142. 


nebidosum, 492. 




Tatusia, 121, 141. 


nuchale, 492. 




hybrida, 462. 


per sonata, 492. 




kappleri, 462. 


sivensis, 492. 




7je/w., 462. 


urahnse, 492. 




Taui-otragus 


woodfordi, 492. 




derbianus gigas, 182. 


Syri-hjiptes 




Taurus 


paradoxus, 519. 




mexicanus, 154. 
Tayassu 
tajacu, 140. 


Tacliybaptes, 78, 82 




Tavra 


fluviatilis, 81, 53l 




barhara, 138. 


Tachyeres 




Taxidea 


cinereus, 514. 




americana, 441. 


Tacliyphonus 




Temenuchus 


coronal us, 474. 




pagodarum, 481. 


melaleucus, 475. 




Testudo 


Tadurna 




elephanfina, 2. 


casarca, 513. 




Tetraceros 


cornuta, 513. 




quadricornis, 455. 


tadornoides, 513. 




Teti-agnatha 


variegata, 513. 




/cTO.r, 297, 302, 303. 


Twuia 




— c?-6ss«, 300, 301. 


le/itocephala, 10. 




Tetrao 


Taeiiioptera 




cupido, 520. 


nengeta, 484. 




j)Iiasia,ncllus. 520. 


Talegalla 




vrogallus, 520. 


lathami, 55, 56, 5: 


23. 


Tetraogallus 


Talpa 




himalayensis, 521. 


■ asiatica, 142. 




Textor 


europcea, 142, 143. 




ali'ctor, 477. 


longirustris, 163. 




Thaunialea 


Tarnandiia 




amhersfi, 6, 7, 56, 


tetradacfyla, 133, 


463. 


621. 


Tainias 




obt.cura, 6, 7, 8. 


asiaticus, 446. 




picta, 6, 7, 8, 5(>, 


lateralis, 446. 




521. 


siriatus, 149, 446. 




Tliaumasia, 313. 


Tamiops 




Tbelutornis 


swinhoei, W9, 384. 




kirtlandii, 3. 


Taiiagra 




Theristicus 


cswa, 474. 




candatus, 510. 


cyan opt era, 174. 




melanups, 510. 


pcdmarnm, 474. 




Therojjithecu.s 


suyaca, 474. 




y^/wrfa, 432. 



xlvlii 



INDEX. 



Tbrasaetiia 

harpyia, 502. 
Tlirasops 

flavigularis, 181. 
Thylaeinus 

cynoce'phalus, 100, 466. 
Tigriosoma 

brasilieiise, 608. 
leucolophum, 508. 
Tinamiis 

soliiarius, 533. 
Tinea, 2l'1. 
Tinnunculus 

alaudariiis, 68, 504. 
cenchris, 504. 
cenchroides, 504. 
domiiiicen»is, 504. 
gracilis, 504. 
QTioluccensis, 504. 
rupicolus, 504. 
f^parverius, 504. 
vesperdnus, 504 . 
Toccus 

erythrorJiynchus. 4-88 
mclanoleiicus, 488. 
Toljpeutes 

quadricinctus, 141. 
tricincius, 141. 
Totanus 

ccdidris, 529. 
fusciis, 529. 
Tracbypbarynx, gen. 
nov., 552. 
nigericB, 552, 554. 
Tragelapbiis 
angasi, 348. 
teko;^^, 348, 349, 350, 

353. 
gratus, 456. 
''scriptus, 2, 456. 
sylvaticus, 456. 
Tragulus 
jauanicus, 3, 460. 
meminna, 460. 
»tanleyanus, 460. 
Tribonyx 

moriieri, 525. 
ventralis, 3, 526. 
Tricbechus 

manatus, 131. 
Tricboglossus 

cyanogrammuii, 494. 
Jhrsteni, 494. 
hfsmatodes, 494. 
mitchelli, 494. 
novcB-hollandim, 494. 
or »a?!MS,' 494. 
ruhritorques, 4, 494. 
Triebosurus 
eanimis, 464. 
J'u/iginos'us, 464. 



Tricbosurus 
orientalis, 464. 
vul'pecida, 464. 
Trigla, 183, 203, 219. 
cora.x, 202. 
gurnardus, 203, 207, 

210, 217, 219. 
7*i)VfWo, 202, 210, 247. 
Tring.a 

alpina, 629. 

canutus, 529. 
Triiigoides 

hypoleucus, 529. 
Triodontopborus, 651. 
Triigon, 60. 
Trypivnosoma 

lewisi, 9. 
Tiipaia 

peguana, 444. 

tewrt, 444. 
Turacus 

6«,ifw^i, 66, 491. 

corythaix, 491. 

l/»mgsto7?i, 491. 

mricrorhynchus, 66. 
491. 

perms, 491. 

schalowi, 491. 
Turdas 

albiventris, 470. 
Turnagra 

crasdrosiris, 472. 
Turnix, 90, 93. 

dussumieri, 524. 

lepurana, 624. 

sykesi, 524. 

taigoor, 524. 

tanki, 624. 

Dar/a, 66, 67, 524. 
Turtur 

aJdahranuf!, 517. 

fkiorqiicdus. 517. 

capicola, 517. 

cardis, 470. 

chinensis, 617. 

commiiifis, 617. 

_ faUdcmdicufi, 470. 

flavipes, 470. 

iliacus, 470. 

leucomelas, 470. 

merula. 470. 

onigratorius, 60, 470. 

musicus, 470. 

viustelinus, 470. 

erientalis, 517. 

poecilopterus, 470. 

pilaris, 470. 

risnrius, 517, 619. 

rufivcntris, 470. 
se?ru/orquafu!<, 517. 
se/icguleiisis, 517. 



Turtur 

siiratensis, 617. 

tigriniis, 617. 

torgwrUus, 470. 

trisfis, 470. 

viaaceiis, 517. 

viscivoriis, 470. 
Tympanistria 

hicolor, 618. 
Tyrannus 

melancholicHS, 484. 

pipiri, 484. 



Uliodon, 297. 

hawkesi. 299, 303, S04. 

(Zora) frcwUum, 306. 
Umbrina, 216. 
Upupa 

epops, 61, 489. 
Uria 

grylle, 532. 
Urobracbya 

albonotata, 476. 

axillaris, 476. 

hocagii, 476. 
Urocissa 

flaviroslris, 483. 

magnirostris, 483. 

occipitalis, 483. 

sinensis, 483. 
Uropsilus, 169. 

sorici2}es, l(i3. 
Ursu.s, 273. 

americamis, 442. 

arei'os, 139, 442. 

— heringia^rus, 3. 

korrihilis, 442. 

isabellhms, 442. 

japonicus, 442. 

7o/or, 140. 

luscus, 139. 

malayaiMS, 442. 

raariti.yius, 442. 

meles, 140. 

ornat'us, 442. 

plscator, 442. 

syriac'iis, 442. 

tlhetaims, 442. 
Urubitinga 

anihraGina, 602. 

meridionalis, 602. 

zonura, .502, 



Vampyrus 

speccrum, 130, 
V-melliis 

cayennensis. 529. 

cr'istaius. 629. 



INDEX. 



xlix 



Vanellus 

vidc/aris. 52'J. 
Vermipsylla, 365. 
Vespertilio 

auritus, 131. 

discolor, 130, 131. 

lejMrinus, 131. 

monpinensis, 162. 

muricola, 162. 

murinus, 130, 131. 

nattcreri, 445. 

noctula, 445. 

■perspicillatus. 130. 

spasma, 131. 

sjpectrum, 130. 

vampyrus, 130. 
Vidua 

alhonotata, 476. 

arclens, 476. 

paradisea, 476. 

principalis, 476. 
Vinago 

calva, 516. 

crassirosh'is, 516. 

delalandii, 516. 
Vipera 

russelli, 4. 
ViveiM-a 

dt'e«a, 137, 438. 

genctta, 137. 

■ichneumon, 136. 

memphitis, 137. 

piitorius, 137. 



Viverra 

tcmgalunya, 438. 

Hbetha, 137, 438. 
ViTerricula 

malaccensis, 438. 

schlegcli, 438. 
Vulpes 

sp., 168. 

a/6a, 135. 

alopex, 134. 

argentatus, 181. 

ccBvulescens, 135. 

famelicus, 1. 

filchneri, 168. 

Aoo^_y, 1(38. 
Vultur 

auricularis, 504. 

calvus, 504. 

monachits, 504. 

occipitalis, 504. 



Xanthocephalus 

icteroceplialus, 481. 
Xanthomyza 

■phrygia, 474. 
Xanthosomus 

jiuvus, 481. 
Xenophrys 

monticola, 393, 397, 
400, 402. 
Xenopsylla 

cheopis, 5. 



Xenorhynchus 

australis, 509. 

senegcdcnsis, 509. 
Xeuurus 

tricinctus, 462. 

unicinctus, 462. 
Xerus 

erythroptos, 446. 

getulus, 446. 

setosus, 446. 



Zapus, 167. 
Zenaida 

amabilis, 517. 

auriculata, 517. 

aurita, 517. 
Zenaidura 

carolincnsis, 517- 
Zeus, 183, 210, 211, 214, 
219. 

ya6er, 209, 216, 248. 
Zonotricbia 

albicollis, 480. 

pileatcc, 480. 
Zorilla 

erythrcBa, 1. 
Zosterops 

dorsaiis, 473. 

lateralis, 473. 

palpebrosus, 473. 

simplex, 473. 



Proc. Zool. Soc— 1911, No. XXXIX 



39 



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LIST OF CONTENTS. 
1911, Paet I. (pp. 1-180). 



EXHIBITIONS AND NOTICES. 

Page 

TfiE Seqretary. Report on the Additions to the Society's Menagerie during the months of 

June, tTuly, August, September, and October, 1910 - 1 

Mr. R. I. PococK, F.L.S., F.Z.S. Exhibitipn of a living specimen of the Black Rat (3/tts 

ratios) from Sar'k 4 

M-r. R. I. PococK, F.L.S., F.Z.S. Exhibition of a female hybrid between a male Black 
Lei^ur {Lemur macaco) and a female oi" the Red-fronted variety of the Fulvous Lemur 
{Lemur futvus rufifrons) •'"> 

The Hon. N. Ciiaules Rothschild, M.A., F.Z.S. Exhibition of some Fleas interesting in 

connection with the spreading of plague by these insects 5 

Mr. J. Lewis Bonitote, M.A., F.L.S., F.Z.S. Exhibition of, and remarks upon, a young Cairo 

Spiny Mouse {Acomys cahiriims) ' ,5 

Mr. J. Lewis Boniiote, M.A., F.L.S., F.Z.S. Exhibition of, and remarks upon, a pair of 
hybrids between the Bramble- Finch {Fringilla onontifringilla) and the Chaffinch {Friii- 
f/illa coelebs) 6 

Mrs. R. Haig Thomas, F.Z.S. Exhibition of, and remarks upon, a series of skins llus- 

trating an experiment in Pheasant-breeding 6 

]>. W. IVrcoLL and Prof. E. A. Mikciiin, M.A., V.P.Z.S. Exhibition of, and remarks upon, 

two species of Cysticercoids from the Rat-flea (Cemz'op^yfes/flsciafKs). (Text-figs.l &2.) 9 

Dr. WifxiAM NicoLL, Demonstration of his method for the collection of Trematodes .... 45 

])i-. R. T. Leiper, F.Z.S. Exhibition of the Nematode infection known as Onchocerciasis. . 45 

* 
Dr. J. F. Gejimill, M.A., D.Sc. Notice of a paper on " The Development of Solaster endoca 

Forbes " 43 

Mr. D. Setii-Sjutii, F.Z.S. Exhibition of, and remarks upon, living examples of the 
.Australian Budgerigar or Undulated Grass-Parrakeet {Melopsittacus tmdulahis), 
showing three colour-phases 45 

The Secretary. Report oii the Additions sto the Society's Menagerie during the month 

of November 1910 100 

Dr. n. Hammond Smith, M.R.C.S., F.Z.S. Exliibition of, and remarks upon, a specimen 
of the Red Grouse, Lagopus scoiicus, which displayed a curious variety of the ordinary 
pluuHige r 100 

Contents continued on page 3 of Wrapper. 



PROCEEDINGS 



GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 



ZOOLOGICAL SOCIETY OF LONDON. 



EXHIBITIONS AND NOTICES. 

November 15, 1910. 

Dr. S. F. Harmer, M.A., F.R.S., Vice-President, 
in the Oliair. 

The Secretary read the following report on the additions made 
to the Society's Menagei-ie during the months of June, July, 
August, September, and October, 1910 : — 

Juxe. 

The registered additions to the Society's Menagerie during 
the month of June were 405 in number. Of these 254 were 
acquired by presentation, 39 by purchase, 18 were received on 
deposit, 12 in exchange, and 82 were born in the Gardens. 

The total number of departures during the month, by death 
and removals, was 142. 

Amongst the additions special attention may be called to the 
following : — 

2 Addra GfizeUes {Gazelki 7'i(Jicollis) d $, new to the Collection, 
1 Mountain Oribi {Ourehia ■moniana), 1 Sabre-horned Oryx (Oj-y.r; 
algazel) $ , 1 Addax [Addax nasomaeulatus), 1 Soudan Fennec 
(Vulpes famelicios), 2 Erythrean Zorillas {^Zorilla erythrcea)^ 
1 Dongolan Genet {Genetta dongolana), and 2 Banded Mongooses 
{Crossarchus fasciatus), from the Soudan, presented by Gilbert 
Blaine, Esq., F.Z.S., on June 4th. 

Proc. ZooL. Soc. — 1911, ISO. I. 1 



2 THL; SECUETAIIV ON' ADDITIOXS TO TUE MENAGERIE. 

1 Acldra Gazelle {Gazella ruficolUs) S , new to the Collection, 
from the Soudan, presented by G. Guy Chetwynd, Esq., F.Z.S., on 
June 4th. 

1 White-spotted Squirrel (Funisciurics leucosiigm.a) and 
1 Speckled Squirrel [Scitirus punctatus), both new to the Collec- 
tion, 1 Gaboon Yiy^ev J^Bitis gahonica), and 1 Nose-horned Viper 
{Bitis 7iasicor7iis), from Dunkwa, Gold Coast, presented by Dr. H. 
G. F. Spurrell on June 18th. 

6 Cocks-of-the-Rock [Rttpicola crocea), from British Guiana, 
deposited on June 14th. 

1 Red Bird-of- Paradise [Pccradisea rubra), presented b}' 
J. M. Doctor, Esq., on June 11th. 

1 Hunstein's Bird-of-Pai'adise (Biphyllodes hunsteini), received 
in exchange on June 23rd. 

1 Short-tailed Pari-ot {Pachynus brack i/ui^us), from the Upper 
Amazons, presented by H. D. Astley, Esq., F.Z.S., on June 14th. 

2 Queen Alexandra's Parrakeets {Sj)athopterus alexandrce), 
from Western Australia, purchased on June 11th. 

2 Condors [Sarcorhamphus gryphus), fi-om Chili, presented by 
the Government of Chili on Jiine 27th. 

A collection of Birds, including Tanagers, Sugax-birds, 
Tinamous, and Conures, from Brazil and Chili, presented by 
Albert Pam, Esq., F.Z.S., on June 18th and 27th. 

A collection of Birds from Venezuela, including various 
Tanag-ers, Finches, a Naked- eyed Pigeon (Columba gymno- 
phthalma), and an Orinoco Goose [Che^ialopex jubaius), presented 
by Hugo Pam, Esq., on June 21st. 

5 Elephantine Toi^toises (Testudo elephantina), from Aldabra, 
presented by Edwa,rd Mayer, Esq., on June 15th. 

July. 

The registered additions to the Societ^-'s IMenagerie duiing the 
month of July were 327 in number. Of these 124 were acquired 
by presentation, 16 by purchase, 85 were received on deposit, 
1 5 in exchange, and 87 were born in the Gardens. 

The total number of departures during the month, by death 
and removals, was 189. 

Amongst the additions special attention may be called to the 
following : — 

1 Bufibn's Kob [Kobus kob), from Sierra Leone, presented by 
J, A. Tinling, Esq., on July 8th. 

1 Sing-Sing Waterbuck {liobtos imcfitostr.s) and two Harnessed 
Bushbucks [Tragelaphtis scripius), from Gambia, presented by 
Capt. Sir George Denton, K.C.M.G., F.Z.S., on July 19th. 

2 Harnessed Bushbucks {Tragelaphus scripttts), I Black-tailed 
Oribi i^Ourehia nigricaudata), and 1 Crowned Hawk-Eagle 
{Spizaetus coronatus), from French Guiana, presented by 
Fenwick Owen, Esq., on July 19th. 

1 Abyssiniaii Duiker (Cephalopkus abyssinicus), from Nigeria, 
presented by W. A. Clayton, Esq., on July 20th. •- 



TIIF. SEORKTARY OX ADDITIOXS TO TIIIO MEXArrERIR. 3 

2 Black-backed Geese (Sarcldiomis melanonota), from India, 
presented by Lt.-Col. D. C. Phillott, on July 25th. 

3 Wattled Peewits {Lohivan^llus lohcttus),?> Bln.ck-tailed Water- 
hens (Trihom/x ventraUs), and 1 TnitedJJmhre {Scopus umhreMa), 
hatched and reared in the Menagerie. 

August. 

The registered additions to the Society's Menagerie during the 
month of August were 200 in number. Of these 105 were 
acquired by presentation, 15 by purchise, 27 were received on 
deposit, 29 in exchange, and 24 were born in the Gardens. 

The total number of departures during the month, by death 
and removals, was 181. 

Amongst the additions special attention may be called to the 
following : — 

3 Grey Orab-eating Dogs (Canis griseus), from the Argentina, 
presented by Wilfred Smithers, Esq., on August 22nd. 

1 Korin Gazelle {Gazella rufifrons) 2 , from Senegal, presented 
by Maxwell Lyte, Esq., on August 2nd. 

2 Ross's Plantain-eaters [Musophaga rossce) and 1 Ardesian 
Hawk [CerchnPAS ardesiacus), both species new to the Collection, 
from Uganda,, presented by L. M. Seth-Smith, Esq., on 
August 15th. 

3 Pigmy Cormorants {Phcdacrocorax pygmceus), from Dobrud- 
sciia, Roumania, presented by the Hon. N". C. Rothschild, F.Z.S., 
on August 25th. 

A collection of Birds, including 2 Raugh-billed Pelicans 
[Pelecanus trachyrhynchus), 2 Roseate Spoonbills (Ajaja rosea), 
and 3 American Darters [Plotus anhinga), received in exchange 
from the Zoological Society of Washington on August 23rd. 

1 Green Tree-Snake (Dendraspis viridis), 1 Kirtland's Tree- 
Snake {Thelotornis kirtlandii), and 1 Banded Gecko (Hemidactyhis 
fasciatus), all new to the Collection, from the Gold Coast, 
presented by Dr. H. G. F. Spurrell on August 21st. 

September. 

The registered additions to the Society's Menagerie during the 
month of September were 228 in number. Of these 37 were 
acquired by presentation, 81 by pxxrchase, 47 were received on 
deposit, 53 in exchange, and 10 were born in the Gardens. 

The total number of departures during the month, by death 
and removals, was 210. 

Amongst the additions special attentio;i may be called to the 
following : — 

1 Brown Bear (Ursus arctos bei-ingiamos), from Kamschatka,, 
presented by W. S. Race, Esq., on Sept. 30th. 

4 Javan Mo\\s,e-Deer (Traguhosjavanic > is), from Java, purchased 
on Sept. 26th. 

J* 



4 MK. K. I. PC)rt»CK ox THE BLACK RAT. 

1 White-naped Roller [Coracias ncevius), from tlie Gambia, 
new to the Collection, deposited on Sept. 15th. 

1 Red-collared Ijovi^eet {Trichoglossusrtihritorques)^hve,(\ in the 
Menagerie. 

1 White-winged Wood-Duck {Asarcornis sactulatus), from 
Burma, new to the Collection, presented by Major H. H. 
Harington, F.Z.S., on Sept. 12th. 

A collection of Snakes containing, amongst others, 6 Indian 
Cobras {Naia tr'qmdians), 2 Banded Kraits (Bungarus fasciatus), 
and 4 Russell's Yipers {Vipera russeUi), received in exchange on 
Sept. 10th. 

October. 

The registered additions to the Society's Menagerie during the 
month of October were 202 in number. Of these 102 were 
acquired by presentation, 26 by purchase, 47 were received on 
deposit, 21 in exchange, and 6 were born in the Gardens. 

The total number of departures during the month, by death 
and removals, was 225. 

Amongst the additions special attention may be called to the 
following : — 

1 Siamang Gibbon (Si/niphcdangus syndactylus), fi'om Sumatra, 
deposited on Oct. 26th. 

1 African Cheetah [Cyncelurus juhatus), from Andamowa, 
German Cameroons, presented by Walter Watts, Esq., on 
Oct. 22nd. 

2 Indian Buffaloes [Buhalus buffehis), from India,, received in 
exchange on Oct. 15th. 

2 Giant Bustards {JSupodotis kori), from P. W. J. Le Roux, Esq., 
a Black-breasted Harrier- Eagle {Circaetus pectoralis), from 
Crossland Robinson, Esq., and a Stanley Crane (Anthropoides 
paradisea), from Dr. Robert Broom, C.M.Z.S., all presented to 
the King's South- African Collection through Dr. L. Peringuey, 
F.Z.S., Secretary of the South-African Pi'esentation Committee, 
and deposited in the Gardens on Oct, 1st. 

A collection of Birds from Venezuela, presented by Hugo 
Pam, Esq., on October 11th, containing a Bare-faced Hangnest 
{Gymnomystax melanicferus), a Chimachima Milvago (Milrago 
chimachitna), a Violaceous Night-Heron {Nycticorax violaceics), 
3 Naked-eyed Pigeons [Columha gymnophthalma)^ and others. 

5 Jackson's Whydah-birds {Drejoanoplectes jacksoni) and 
2 Crimson-ringed Wh3'dah-birds {Penthetria laticauda), from 
East Africa, new to the Collection, presented by Mrs. George Style 
on Oct. 22nd. 

Mr. R. I. PococK, F.L.S., F.Z.S., exhibited a living specimen 
of the Black Rat [Mas rattus) from Sark, which had recently 
been presented to the Society's Menagerie by Mrs. C. Russell. 



U\ XKWI.V-HUHX CAJKO Sl'lNV MICK. 

Mv. R. I. PocoCK ulso exhibited a female hybrid, bred in the 
Society's Gardens, between a male Black Lemur {Leimor macaco) 
and a female of the Red- fronted variety of the Fulvous Lemur 
(Lemur fidvus rujifrons), and pointed out that the offspring re- 
sembled neither of its parents. It had not inherited the facial 
fringe of its father nor the white over the eyes and on the fore- 
head of its mother, this ai-en. of the head being dark ashy black 
witli a deeper tinted central line, more approaching the colour 
seen in the Black-fronted variety of the Fulvous Lemur (Z. fidvus 
nigrifroiis). 

The Hon. N. Cpiarles Rothschild, M.A., F.Z.S., exhibited the 
following species of Fleas, which were of interest in connection 
with the spreading of plague by these insects : — Pidex irritans, 
Xenopsylla cheo^iis, Ctenocejihalics canis, 0. fells, Cerato2)hyllus 
fasciatus, C. londimensis, Ctenopsylla mai/ardi. 



Mr. J. Lewis Boxhote, M.A., F.L.S., F.Z.S., exhibited in 
spirit a young Caii'O iSpiny Mouse {Acomys cahirinus) about 12 
hours old, and pointed out the advanced state of the young a,t 
birth as compared with the common House-Mouse and other 
species of the genus Jlas. 

In Acomys the 3'oung at birth Avei'e of a pale slate-colour and 
sparsely covered on the upper parts with downy hairs 2 to 3 nnu. 
in length. Across the lower j^art of the back the spines could 
be distinctly felt. 

In spite of this advanced development the period of gestation 
was only 1 1 days at the most. In the case under notice the male 
was introduced at 7 p.m. on 19th July and the young were born 
between midnight and 9 a.m. on the 31st July, a maximum 
period of 11 days and 14 hours. The period in the House-Mouse 
was about 13 days. In Acomys the usual number in a litter was 
three ; four were exceptional, but two were not uncommon. 

In correlation wdth their forward development the j^oung were 
also very large at birth, as a comparison of measurements of the 
individual exhibited and the average measurements of an adult 
male show : — • 

Young 
12 hours old. Ad. (? . 

Head and body ... 51 mm. 101 mm. about | full size. 

Tail 35 mm. 105 mm. ,, g ,, ,, 

Hind foot 14 mm. 18 mm. ,, | ,, ,, 

Ear 8 mm. 17 mm. ,, g ,, ,, 

When 8 to 10 days old the young were able to take care 
of themselves, and on the fourth day after birth their eyes were 
open and they occasionally left the uest and ran about the cage. 

They bred at a very early age, and by the agency of a young male 
born on the 4th ,Tune progeny were produced on the 31st July, 
so that it must have bred Avhen only 46 days old. 



MRS. R. HAIG THOMAS OX 



Mr. Bonhote further stated that among Sand-Rats of the genus 
Meriones, a considerable number of which he had bred this year, 
the period of gestation was at least 17 days, and the young, which 
were born quite naked, looked after themselves in about three 
weeks, five being the usual number in a litter. 



Mr. J. Lewis Bonhote, M.A., P.L.S., F.Z.S., exhibited a pair of 
hybrids between the Bramble-Finch {Fringilla montifriiigilla) and 
the Chaffinch {Fringilla coelebs). This cross had been bred for the 
first time in 1907 by a lady in Kent, who had since bred several 
every year. 

Mr. Allen Silver, a well-known fancier, first called Mr. Bonhote's 
attention to these birds. He wrote : — " The young in nestling 
plumage chiefly resemble a nestling Chaffinch, but show traces of 
the white rump. 

" Males from the cross in which the Brambling was the cock 
are in every case more brilliant and richer in tone, and exhibit a 
marked Bramble-Finch form of head, more so than the cross in 
which the Chaffinch was the male parent. The heads of the 
males in this latter cross (Chaffinch J and Brambling $ ) are less 
speckled and almost even in tone of colour, showing few signs of 
the dark bases to the feathers on the head and having more of a 
Chaffinch shape." 

The cock exhibited resembled a, Bra-mble-Finch on the head, 
rump, tail, and vent, but on the mantle and breast the colour 
showed a mixture of the two species; the tail also shoAved more 
white than was usually found in a pure-bred Bramble-Finch. 

A peculiar feature of this bird was a slight " peak " crest, which 
had not appeai^ed in any of the other specimens. It seemed to be 
caused by the long feathers of the nape (a Bramble-Finch feature) 
meeting the long feathers of the crown (a Chaffinch feature). 

One other male of this cross that Mr. Bonhote had seen closely 
resembled a hen Bramble- Finch. 

The hen bird exhibited showed the Chaffinch blood more 
clearly, her only Bramble-Finch features being the two parallel 
stripes on the nape, the partially white rump, traces of j^ellow on 
the wing-bars, and a more yellowish tint on the breast. The 
fertility of the hybrids had not yet been tested. 



Mrs. R. Haig Thomas, F.Z.S., exhibited a series of skins illus- 
trating an experiment in Pheasant - breeding, and gave the 
following account of the production of Thaumalea ohscm-a in the 
F, generation from a cross between Thaumalea amhersti $ and 
T. picta S '■ — 

" The skins of the birds used in this experiitient are exhibited 
together with their pedigree (see page 7). An Amherst cock is 
placed with these skins to show the plumage, but the bird' took 
no part in the experiment. 



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5 ON EXrERIMKNTAL PHJBASAXT-BKBEDING. 

" In Elliot's ' Phasianidse ' is a coloured plate of a pair of 
Thaumalea ohscura with their young, which were considered by 
him and some other ornithologists as a variety of picta ; according 
to this plate the birds apparently bred true. 

"In my pheasantry in 1907, three Amherst hens were mated 
with a Golden cock and produced a number of young called F, in 
the pedigree. In 1909 two pairs of these F, birds were mated, 
and from these two pens 20 F^ chicks were hatched. Amongst 
them were three chicks of a deep chocolate-brown, in startling 
contrast to the others, which were cream-colour with a bronzing of 
russet on the back and throat. The down of these ohscura chicks 
Avas of a uniform dark brown to the skin, and each eye was 
rimmed with a, finely pencilled cream line, which was connected 
by another cream line across the top of the bill, giving the quaint 
appearance of a pair of specta,cles ; there was also a cream patch 
on the throat, varying in size in each individual, sometimes being 
a mere spot. The legs were a dull olive-brown. These brown 
chicks were produced from both pens, and were of both sexes, 
but unfortiuiately only one of the three (a cock) was reared. 
Thamnalea birds attain adult plumage only in the second }'ear, so 
it was not till July this year that my surmise that ohscura 
had been produced was confirmed. The cock has the same barred 
tail (pattern Amherst, coloration C4olden), and the same dai-k 
brown on head and throat and neck, as the bird depicted on Elliot's 
plate, but his breast is a duskier red and the crest and mantle 
paler. When a chick, his coloration was similar to the young 
shown on Elliot's plate, only the brown was a deeper, richer shade. 
This F^ cock was mated in the spring of 1910 with two F^ hens, 
his sisters. The darkest hens were picked out, with Golden eye- 
skin (yellow with red round the rim) ; Amhei'st hens are lighter 
and greyer in plumage than the Golden, and the eye-skin is a 
greenish blue. From this mating eight birds were hatched, four 
cream-coloui-ed and four dark brown chicks ; these last had pre- 
cisely the same coloration as that of the father. The numbers 
tally with those to be expected according to Mendel's laAv. I 
hope to obtain more evidence next season from the same birds, 
and to have better luck, for all these eight chicks lived only a few 
days. As I intend to breed again from him, I caimot show you 
the skin of the a,dult ohscura cock, biit only some feathers plucked 
from his breast and back, and the skins of his two brothers and 
of two of his sisters. An examination of these breast-feathers 
plucked from ohscura and of some of the breast-feathers of the 
cock marked '■ s,eYa\- ohscura' will show the reason of the so-called 
' duskiness' of hue ; each feather has a bar of metallic green on 
the inner half, while the outer end of it is led. You will observe 
in the Golden cock that the breast-feather is red throughout. 

" Apparently, then, Thaumalea ohscura is a hybrid recessive 
mutation, breeding true, as shewn in Elliot's plate and also in my 
experiment this year, according to Mendel's law. I ha^e used 
the word ' mutation,' but these birds may possibly be a I'eversion 
to the ancestral parent form of both '' species" of Thamiialea. 



ON' TWO SPKCIKS OF CVSTICEilOOIDS. 9 

" By whatever name it is called, this hybiid is undoubtedly a 
homozygote for pattern and coloni-, pine and permanent, trans- 
mitting these characters to its descendants. Cross-breeding 
between these two varieties of Thamnalea pi'oduces a new form, 
owing possibly to the meeting of characters never previously com- 
bined, and as they are constant it is evident these have an affinity 
and have become insepai-able. 

" On such lines Evolution might be conceived as having pro- 
ceeded fairly rapidly towards the sepai'ation of species. 

" Some while ago 1 showed a hybrid hen whose pure Swiiihoe 
plumage had been transmitted through the cock. To-day the 
skin of her son, a pure iSwinhoe cock, is exhibited for your 
inspection. 

'' An account of my method and management of these Pheasant- 
breeding experiments has been already published in the ' Pro- 
ceedings ' (1909, p. 885), showing the care and precautions used." 



Two iSpedes of C ysticercoids from the Rat-flea (Ceratophyllu.s 
fasciatus). 

(Text-figures 1 & 2.) 

Dr. W. NicoLL and Professor E. A. Minchix, M.A., Y.P.Z.S., 

exhibited examples of two species of Cysticercoids which had 
been found in the body-cavity of the i-at-flea {Cerato2yhylh(,s 
fasciatus) dissected by Professor Minchin in the course of in- 
vestigations upon the development of Trypanoso'ma leioisi in the 
flea. 

The first specimen shown (text-fig. 1) was one previously ex- 
hibited by Piof. Minchin (P. Z. S, 1909, p. 741). >Since "then 
experiments had been carried on by Dr. Nicoll at the Lister 
Institute, which proved that it was the larval form of Hymenolepis 
dimiiiuia, and threw light upon the mode of infection. This 
Cysticercoid had occurred in about four per cent. (8 in 207) of all 
the fleas examined during a period of thirteen months. It was 
found in the body-cavity and usually only one specimen at a 
time. As many as three, however, had been found in one flea. 

In the reti'acted state the Cysticercoid consisted of a flattened 
oval body and a moi-e or less elongated tail. The dimensions of the 
body were '31 x '25 mm., and the tail might be as much as "8 mm. 
long. The wall of the body consisted of a number of well- 
difierentiated layeis. The outermost layer was non-cellular and 
bore a cei'tain resemblance to cuticle. It was marked by fine 
radial striations. The second layer consisted of columnar cells, 
each with a large and distinct nucleus. ISTeither of these layers 
occurred in the tail. The third layer w-as parenchymatous, 
consisting of a large number of irregularh' disposed cells. This 
was continuous with the substance of the tail. The next layer 
was apparently of a fibrous nature, consisting of a few cells set 
in a mass of circularly arianged fibres. This layer was sejmrated 
from the next by an intervening space. This was seen usually 



10 



DR. W jSJIOOLL and PROF. E. A. MIXCHIN OJT 



only as a narrow fissui^e. It represented the very much reduced 
vesicle, and separated the main part of the body from the part 
which was eventually evaginated to form the head of the tape- 
worm. Internal to this was another layer of loose parenchyma, 
which at the posterior pole merged into the tissue of the scolex. 
The latter was extremely small (-075 x -09 mm.). On it there were 
four small circul;a- suckers (diam. "OSS mm.) and a small unarmed 
rostellum. 

■ Text-fig. 1. 











Cysticercoid of Si/menolepis diminuta. X 125. 

This description tallied almost exactly with that of Gi'assi and 
Rovelli (Atti Ace. Gioenia Sc. Nat. Catania, iv. 1892, pp. 31-33, 
pi. iv. fig. 3). They found a corresponding Cysticercoid in the 
beetles J-X-is spwiosa and Sccmrus siriatus, in Anisolabis anrmlipes 
and in the larva of Asopia farinalis. By feeding experiments 
with Cysticercoids derived from the first of these, they were able 
to produce infection in a man, and the tapeworm which was 
recovered wsiii Bymeoiolepis diminuta Rud.( = TcEm« leptocephala 
Creplin). 

It was recognized that the Cysticercoid from the rat-flea wa& 
very probably the larva of Hymenolepis dimimita, but that could 
not be definitely stated without further information, for two 
other rat -tapeworms {H. relicta Zschokke and IT. horrida 
v. Linstow) are known, closely resembling H. diminuta and 
probably possessing larvse almost indistinguishable from that of 
H. dimirncta. On that account, in order to establish the identity 
of the Cysticercoid, a series of feeding experiments was undertaken. 



TWO Si'liClES OF CYSTICKRCalDS. 11 

For tliis purpose a litter of young rats was separated froui tlieir 
mother as early as possible and isolated. They were fed entirely 
on boiled bread and milk. They were kept under observation 
for three months, during which time frequent examination of 
their faeces showed that they were free from tapeworms. Two of 
the rats were then fed daily with the fleas, which were mashed up 
in their food. This was continued for 19 days, feeding being 
omitted on three of these, when one of the rats died. During 
this period the faeces were examined regularly, but no ova were 
found. On examining the dead rat, five specimens of Hymenolejns 
dlminuta were found in the intestine. They wei'e of various 
sizes, the largest being over 40 cm. long and the smallest only 
1 cm. Next day numerous ova were found in the fasces of the 
other rat. From this it was evident that the complete develop- 
ment of Hymenoleijis diminuta took place in less than three 
weeks. Grassi and Rovelli found ova after 15 days, which was 
probably about the actual length of time required. The remaining 
rats of the litter were kept under observation for a further period 
of two months, but during that time no ova were found in the 
faeces of any of them. The same experiment was repeated later 
with a similar result, ova appearing in the faeces on the 19th day. 
The rat which survived the first experiment was eventually 
killed and nine tapeworms were removed from its intestine. A 
total of 14 tapeworms was therefore pi'oduced from the 340 fleas 
with which the rats had been fed. This indicates that about 
four per cent, of the fleas contained Cysticercoids, and the result 
agrees with the number of Cysticercoids found in fleas which were 
actually dissected. 

The supposition, put forward in the previous communication, 
that the flea becomes infected in its larval state, had to some 
extent been confirmed. The intact ova of Hyvienolepis diminuta 
were found in the intestines of 2 larvae out of 145 examined. No 
stages in the development of the Cysticei"coid, however, had been 
yet met with in the larval flea. On the other hand, a very young 
Cysticercoid had been found in the body-cavity of one out of a 
dozen pupae examined. The presumption was that it was only 
the oldest and largest flea-larvas that were capable of ingesting 
the tapeworm eggs and that no development took place until the 
beginning of the pupal stage. 

The second Cysticercoid (text-flg. 2, p. 12) was of much greater 
interest than the first. Hitherto only a single specimen had 
been met with, and it occurred in the body- cavity of a male 
Cercdophyllus fasciatus. It was considerably smaller than the 
first Cysticercoid, and like it consisted of an oval body and a tail. 
The former was "16 mm. long and the latter '19 mm. The two 
outermost layers of the body-wall were the same as before, 
but the parenchjmiatous and fibrous layers were not so sharply 
differentiated from each other. The scolex, again, was very much 
larger (diam. '096 mm.) and it had a rostellum armed with a 
single row of spines, OS' in number. The length of each spine was 



12 



BK. W. NICOLL AND PKOF. E. A. MIXCUIN ON 



•017 mm., the handle being "01 mm. and the guard '007 mm. 
long. The prong was of the same length as the guard. There 
were four well-formed suckei^s, which had been pressed into a 
somewhat crescentic shape (greatest diameter "042 mm.). 



Text-fiir. 2. 








V ■*./ 

CysticevcoiA oi H^menolepis mi(ri)ia. X 250. 

a. Rostellar spine in full lateral view. X 600. 

b. The same, somewhat tilted. 

Besides the two Cysticercoids described here and the Avell-known 
Oysticercoid of Dipylidium caninum from the dog-flea and humaii 
flea, there was only one other instance on record of a Oysticercoid 
being found in a flea. This observation was made a few months 
ago by Dampf (Oentralbl. f. Bakt. etc. Ite Abth. Orig. liv. 
pp. 452-4), who found in a new species of flea Mesopsylla evxita from 
the jerboa (Alactaga jaculus) a Oysticercoid bearing a close resem- 
blance to the second of those described here. Dampf was obliged 
to make his examination from a mounted (type) specimen of the 
flea, and on that account could not be certain of some of the 
details, as, for instance, the exact number of the spines and the 
presence of a tail. So far as he could make out, there were from 



TWO SPECIKS OF CVSTICERC'OIDS. 1 ."{ 

20 to 24 spines. Llilie, to wliom tlie specimen was refevi'ed, 
mentioned the supposition tliat the Cysticercoid might be that of 
Ilyinenolepis nana (v. vSieb.). 

From the description given of the second Cysticercoid it was 
apparent that it was very like the scolex of Hymenolepis mur-ina 
I)uj., a freqnent parasite of vaiious species of rats. Tlie size of 
the scolex of the Cysticercoid, the arrangement, number, size, and 
shape of the spines all agreed with those of Hymenolepis murina, 
and it was certainly the only rat-tapeworm to which the Cysti- 
cercoid could be referred. Tlie closely allied Hymenolepis iniet'o- 
stoma Duj. had 30 spines of much smaller size, //. contracta 
Janicki and H. maris variegati Janicki, neither of which corre- 
sponded to the Cysticercoid, were the only other ai-med Hymeno- 
lepids found in rats, ami the authors were forced to the conclusion 
that this C3^sticercoid must either be I'eferred to H. 7nurina or 
to some other as yet vmdescribed form, the scolex of which was 
indistinguishable from that of H. mu?-ina. 

Hitherto the life-history of H, viurina had been believed to be 
of a unique type. According to the researches of Grassi and 
Rovelli {op. cit. pp. 75-83) this tapeworm dispensed altogether 
with an intermediate host, and they had demonstrated that it 
passed its encysted stage in the walls of the intestine of the rat 
itself. This was supported on general gi-ounds by the fact that 
this particular tapeworm usually occuired in enormous numbeivs, 
which would involve the ingestion of a correspondingly large 
number of intermediate hosts. The matter gained interest from 
the fact that H. murina was believed by some to be identical with 
H. nana, a dangerous tapeworm of man, although the identity was 
still a matter of dispute. In size and structure the two species 
w^re indistinguishable, and H. nana like //. mnrina occiu-red in 
enormous numbers. Further, no intermediate host had yet been 
discovered for H. nana. Now the discoveiy in the rat-flea of this 
Cysticercoid bearing such a close resemblance to the scolex of 
H. 'murina led to one of two conclusions : either that, as already 
mentioned, there might be some undescribed tapeworm of which 
the Cysticercoid was the larval stage, or that the rat-flea might 
function as an intermediate host of H. murina. The latter 
supposition, even if coirect, did not necessarily disprove Grassi and 
Rovelli's results. Rats certainly ingested a large number of fleas 
from time to time, but it was difficult to imagine that they would 
acquire a very large infection with H. mu7^ina by this means ; 
and if H. nana be identical with H. murina it was scarcely 
conceivable that a human being could accidentally swallow a 
sufficient number of fleas to give rise to an infection of over a 
thousand tapeworms, as was frequently found to be the case. It 
would still therefore be necessary to admit that the development 
of H. marrna usually took place without an intermediate host, 
but that occasionall}^ the rat-flea might function as such. With 
such a conclusion the life-history of H. murina became even 
more remarkable than before. 



u 



MR. J. LEWIS BOXHOTE OX THE 



PAPERS. 

1. On the Inheritance o£ the Webfoot Character in Pigeons. 
By J. Lewis Bonhote, M.A., F.L.S., F.Z.S. 

[Received August 29, 1910: Read November 15, 1910.1 

(Text-figures 3 & 4.) 

In the P.Z.S. for 1905, p. 550, Mr. Staples Browne published 
a paper on the webbed-foot character in Pigeons, and from a 
series of careful experiments conducted by him the results seemed 
to show fairly conclusively that the webbed foot was a Mendelian 
character and was recessive to the normal or non-webbed foot. 

In this paper the author drew an arbitrary line and counted as 
webbed all birds which showed on one foot a web at least to the 
first interphalangeal joint of the 2nd and 3rd digits and to the 
second interphalangeal joint of the 4th digit, any bird with 
less webbing being considered as normal. In my experiments 
I have followed his distinction for the sake of uniformity. All 
his results, with one exception, go to show that the webbed foot 
behaves as a pure Mendelian recessive chai-acter. 

In the aberrant case tw^o F^ birds were mated together for two 
seasons, producing altogether 23 birds, in none of which the web 
appeared. Both these F^ birds were subsequently tested and proved 
to be carrying the webbed character. 

In thespi'ing of 1905, Mr. Staples Brow-ne kindly gave me a 
pair of his webbed birds [Exp. 14, toe. cit.], and during that year 
and the two subsequent years this pair produced nothing but 
webbed birds. 

In 1907 Mr. F. W. Smalley, a well-known and successful breeder 
of fancy Pigeons, w^rote to me in regard to some webbed birds 
which tended to make their appearance from time to time in his 

Table I. , 



Exp. 
No. 


?• 


Origin 
from 


Also 
used in 


(?. 


Origin 
from 


Also 
used in 


Nature of 
Mating. 


Fe 

Normal. 


et. 
Webbed. 


a ... 


N (61) 




c 


N (1'26) 


h 




DRXDR 


6 


2 


h ... 


N(29) 






N (126) 


a 




D XDR 


6 





c ... 


N(61) 




a 


N (104) 






DRXDR 


6 


2 


d ... 


N (10) 


h 




N (01) 


c 




DRXDR 


6 


1 



stud. At my request he kindly sent me their pedigrees, from 
w^hich it will be seen. that the inheritance was strictly Mendeliaii. 
and entirely bore out Mr. Staples Biwvne's results. . 



WEBl'UUT CHARACTER IX PIGEUXS. 

Text-fig. 3. 
Nc3xN$ N6x N9 



15 




JOTHERSN 2W 



No JotHERsN 



W69 



Pedigree of Mr. Smalley's bird (W 69). 
(Birds underlined ai"e supposed lieterozygotes.) 

A fui'ther and striking confirmation of this came to light this 
year (1910), when a web-footed bird suddenly appeared in whose 
direct ancestry on either side no webbed birds had been known 
for at least six generations ; an investigation of the pedigree, 
however, showed one of the birds to have been a possible hetero- 
zygote, and on this assumption, and presuming all the subsequent 
matings to have been DR x D, the case is quite simple. To some 

Table II. 



Exp. 

No. 


' ?. 


Origin 
from 


Also 
used in 


$. 


Origin 
from 


Also 
used in 


Nature of 
Mating. 


Fe 
Normal. 


et. 
Webbed. 


e ... 


N(5) 


a 




N"(50) 






DRXD 


3 





f - 


N(52) 


e 




N (30) 






DRXD 


7 





ff ... 


N (72) 


f 




N(ll) 






DRxD 


4 





h ... 


N(46) 






N(d4) 


9 




D XDR 


15 





i 


N (1376) 






N (1354) 


h 




D XDR 


13 





k ... 


N (5486) 


i 


... 


N (4863) 


h 




DRXDR 


2 


1 



it may appear strange that a DR bird should have been chosen 
in so many successive years, when the chances were even on the 
bird being a DR ,or pure jSTormal. . It must, however, be re- 
membered that since young were. I'eared from a lai-ge proportion 



16 



MR. J. LEWIS BOXIIOTE OX THE 



of the birds noted in Table II., one would, in theory, have 
expected the web to have made its appearance in more than one 
instance, as the number of DR birds in Mr. Smalley's lofts must 
be considerable. 

The fact i-emains that a webbed bird has been bred, and if its 
pedigree were not well-known one would have liad to regard it 
as a " sport," without being able to accoiuit for its presence in 
any way. 

So far, therefore, from well-attested evidence of independent 
breeders, the webbed character seems to be established beyond 
doubt as a Mendelian recessive. 

In the summer of 1907, however, Mr. Smalley sent me a male 
web-footed bird (W 69), which I paired at once with one of the 
progeny (also webbed) of the pair of birds originally given me by 
Mr. Staples Browne. My object was merely to get a change 
of blood and to keep up a small stock of webbed birds. 

Both the parents were webbed and, .as we have seen, their 
ancestry showed that in both families the web had acted as a, 
Mendelian recessive, so no doubt was felt but that the whole of 
the progeny would also be webbed. The result of this mating 
was five birds, of loMchfour tvere normal. 

For the past three years I have carefully mated these five 
birds and their descendants, with the results as shown in the 
follovving table : — 

Table III. 



Exp. 

No. 




Origin 
from 


Also 




Origin 
from 


Also Nature 


Feet. 




?• 


used 


(?. 


used of 


Normal. 


Webbed 










in 






in 1 Mating. 








1 .. 


W (7) 


14 SB 




W(69) 


d 




RXR 


4 


ll 


Mixed 

strains. 


2 


W (a) 


1 


14 


N (d) 


1 


:::-! 




1 


3 '.'. 


N (b) 


1 


4 


N (13) 


1 


13 




1 


1 




4 .. 


N (b) 


1 


3 


N {<■) 


1 






5 




Mixed 

strains. 


5 .. 


W (24) 


2 


12 


W (9) 


14 SB 






1 


6 .. 


N (29) 


4 




W (35) 


2 




in 


5 


7 .. 


N (22) 


4 




N (19) 


4 




1 theory 


9 


1 




8 .. 


W(23) 


4 




W (20) 


4 


.. 


' ' all these 





2 




9 .. 


W (27) 


3 




N (543) 


3 




matings 


2 


2 




10 .. 


N(533) 


7 




w (36) : 2 




must be 





ii 


Mixed 
strains. 


11 .. 


W (14) 


USB 




W (30) i 4 


12 


1 RxR! 


1 


12 .. 


W (24) 


2 


"5 


W (30) ! 4 


11 


j 


4 


13 .. 


W (26) 


11 




N (13) 1 


2 




3 


»{ 


Mixed 


14 .. 


W (a) 


1 


2 


W (8) 14 SB 


.J 




2* 


strains. 



* In both these birds the amount of webbing is just on the line fixed by 
Mr. Staples Browne. 



If we look closely into the foregoing table, we may note that 
whenever the apparent Normals are mated together we get some 
webs. This has been done in experiments 3, 4, and 7, the total 
result being 12 normals to 6 webs. Were these normals hetero- 
zygotes Ave should have expected 12 Normals and 4 Webs. 



WEBFOOT CHARACTER IX PIQEOXS. 



17 




Proc. Zool. Soc— 1911, No. II. 



18 ON THE WECFOOT CHARACTER IN PIGEONS, 

If we take the ISToi-mal-Web matings, as in experiments 2,6, 
9, 10, and 13, we find the total results to be 11 Normals to 
10 Webs. In this case were the Normals heterozygotes we 
should expect equality, which is practically the result attained. 

Lastly, when we mate Webs to Webs, as has been done in 
Exps. 1, 5, 8, 11, 12, and 14, we should expect Webs only, and 
this result was only attained in one case (Exp. 8), where only two 
birds both webbed were reared. In the other five cases together 
15 birds were reared, 12 Normal and 3 Webbed. The proportions 
are almost exact for a DR x DR mating, which, of course, we 
know they cannot be ; but it is not without significance to note 
tha,t in four out of these five experiments the matings Avere a 
cross between the mixed strain and Mr. Staples Browne's strain. 

These, then, are the facts, and at present it seems difficult 
to reconcile them with the Mendelian theory, although they 
certainly seem to show that the inheritance of this character is 
to a cei'tain extent in a,ccoi-dance with that theory. 

According to previous and contemporary work on this character, 
no individual should have had a normal foot. If, however, the 
Normals that have appeared be considered as heterozygotes, then, 
considering the small numbers, the results from the matings are 
not greatly at variance with what we should expect. 

As regards the Webs (in these experiments), when mated with 
Normals they behave as recessives, but when mated with other 
Webs of either the mixed or of Staples Browne's strain they 
produce Normals to Webs in a ratio closely approximating 3:1. 
Such a result in the case of no less than five matings seems to 
show pretty conclusively that the joining of the two strains has 
produced a factor disturbing the normal course of the Mendelian 
inheritance. 

It must be carefully noted that each strain by itself breeds 
true according to Mendel's laws and that the disturbing factor is 
not brought in from one side only, but is due to the intermixture 
of the two strains. 

Mention should perhaps be made of a suggestion by Prof. Bate- 
son, to whom I wrote on the subject. This was that the webbed 
foot was possibly a double character and that the web between 
digits 2 and 3 had a separate inheritance from that between digits 
3 and 4. Now, if we look at Exp. 1 in this light, we find that 
in I the male is very slightly webbed (practically normal) and 
that in | he is fully webbed. In the female, on the other hand, 
the webbing in | is very full and in | extremely small. 

We could thus suppose the male in Exp. 1 to be | N (W) 
I WW, female § WW | N (W) ; this would then give in F, 
^ Noi'mals and Webs equally, | Normals and Webs equally, so 
that we might expect pure Normals (| N | W, | W | N) and pure 
Webs in equal numbers — as a matter of fact, 4 were Normals and 
IfNfW. 

This suggestion, however, becomes untenable for other reasons. 
If we examine the birds I bred for Mr. Staples Browne (Exp. 14, 



ON THE CRANIAL CHARACTERS OF LACERTA JACKSONI. 19 

loG. cit.), together with 5 more individuals reared since, we find 
that the old male no. 19 SB was undoubtedly | WW | N (W) and 
the female was the same ; consequently all birds bred from this pair 
should be webbed in | and a quarter of them webbed in |. As a 
matter of fact, out of 10 birds, 1 is normal in |, 3 are webbed 
in |, and one has all fonr toes f idly loehhed. 

It is unnecessary, therefore, to follow up the matter further, as 
slight deviations appear in almost every mating. 

We must, therefore, be driven back to the suggestion that 
a factor composed of two parts (cryptomeres), one of which is 
contained in each strain, causes the web to be suppressed or 
concealed. 

On this assumption none of the results attained are at variance 
with the Mendelian theory. The proportions, however, do not 
fit in, but as the numbers are small this does not count for much. 
On the average the extent of the webbing shows a tendency to 
diminish and the foot to approximate to the normal as compared 
with birds of either of the pure strains, and in only one bird, bred 
in Exp. 9, has the web included the hallux. 

Owing to lack of space and pressure of other experiments, I 
do not propose to pursue this inquiry further, but should anyone 
wish to follow it up I shall be very pleased to place all my material 
at his disposal. 

My best thanks are due to Mr. Smalley, who has not onl}'- 
supplied me with full and accurate particulars from his pedigree 
book, but has also during the past year kept and bred many of 
the birds in his lofts. 



2. Notes on the little-known Lizard Lacerta jacJcsoyii Blgr., 
with Special Reference to its Cranial Characters. By 
Edward Degen, F.Z.S, 

[Received September 15, 1910: Read November 15, 1910.] 
(Text-figures 5-7.) 

Through the kindness of Mr.G. A. Boulenger, F.R.S., V.P.Z.S., 
a series of seven specimens of Lacerta jacksoni has been placed 
in my hands for investigation. These specimens, recently pro- 
cured by Mr. R, Kemp at Mumias, Mount Elgon District, 
British East Africa, at an altitude of 7000 feet, reached the 
British Museum in a bad state of preservation ; opportunity was 
therefore taken to submit the cranial structure of this little- 
known species to a careful examination, the results of which are 
now placed on record. 

Of the seven specimens six were males and one female. 

The specimens from which the skulls were prepared were a 
small, but quite adult male, measuring 70 mm. from snout to 
vent, and the single female, measui-ing 68 mm. The dimensions 

2* 



20 



MR. EDWARD DEGEN OX THE 



of the former tliei-efore are about the same as those of the single 
male specimen from which the species was originally described 
and figured*, after its discovery at "Ravine Station," Man 
Mountains, British East Africa, at an altitude similar to that at 
■which the specimens now studied were obtained. 

Description of the Skull of the Male. 

Dimensions, — Total length 21 mm. ; width 12 mm. (as measured 
between the extreme points of the zygomatic processes of the 
jugular bones) ; height 6 mm. 

Its width is thei^efore contained 1-9 times in its length, and its 
height 3-3 times. 

Text-fiG-. 5. 






'^^'^«^,;^ 




Skull nf male. 
Tipper, lower, lateral and posterior aspects. X 2^. 

General Configuration. — Remarknbly depressed, as may be 
noted from its proportionate dimensions, and fuither seen in the 
above figure representing the profile. 

Facial portion. Snout short, broadish at base, terminating in 
a- bluntly rounded-off prsemaxillary. 

* " On New Lizards from the Interior of British East Africa," by G. A, Bo\ileng-er) 
F.K.S., Proc. Zool. See. 1899, pp. 96 & 07, ]il. x. 



CRAXIAL CnAUACTERS OF LACERTA JACKSONI. 21 

Craxium [Fronto-pa7'ietal portion). — Squai-ish. The whole of 
the upper surface from the anterior borders of the nasals to the 
posterior margin of the pai-ietal quite flat, and forming an almost 
perfectly level pla,ne. 

Denned incrustation. Evenly spread, profuse, but highly 
porous, and covering the whole of the upper surface, including a 
small portion of the nasal process of the prtemaxillary l)one. 

Lamina supraciliaris (text-fig. 6, A, p. 23) (Supra -orbital 
region). Completely ossified ; 4 supraoculars, and 1 supraciliary 
on each side. 

Supraorbital hones (text-fig. 6, B). Small ; irregularly tria.n- 
gular and thickish, convex laterally. The internal angles termi- 
nating in a hook-like spine and the posterior margins irregularly 
serrated. Their lateral edges barely showing on the exterior and 
almost wholly concealed under the first supraciliary scutes. 

Pteri/yoicl bones. Toothless ; moderately diverging immediately 
in front of the dilated lateral processes of the basisphenoid. 

Postfrontals forming a, single plate. 

A Retrociliary present (text-fig. 6, A). 

Dermal ossification of Temporal region. Supratemporalia 1 and 
2 partially ossified. 

Dentition. Number of teeth in preemaxillary 9 ; in each 
maxillaiy 18. 

Nasal apertures (text-fig. 6, D). Subcircular, the longitudinal 
diameter the greater. 

Parietal Region. — This, as in all Lacertidse, is clearly defined 
in contour, and forms the greatest expanse of the cranial roof. 

Its anterior borders practically are coincident with the fronto- 
parietal suture and the postorbital margins of the adjacent post- 
frontals. Posteriorly, the parietal is produced well ba,ckwards, 
and ends in a straight edge, the latter being the result of an 
increased deposit of the crusta calcarea — a feature mostly absent 
in the very young, in which this portion is concavely crescentic. 
This edge connects the two outward and backwardly directed 
processus parietales, spanning over the parotic processes, the 
processus ascendens of the supraoccipital bone, and covering the 
greater part of the latter in this species, so that the pars condy- 
loidea of the occipital alone projects beyond it when the skull is 
viewed fi'om above. 

The parietal processes, slightly triangular in a transverse 
section at their terminal parts, are quite flat at their basal 
portions and are mere continuations of the median portions of 
the parietal itself. They have not, as is the case with the 
majority of the Wall-Lizards in which the facies cranialis is more 
elevated, the downward!}^ directed ridges set at right angles with 
their bases, in order to establish the contact with the underlying 
supraoccipital. 

Jiio Foramen pariefale. (For special remarks on this striking 
feature refer to p, 29 at the end of the description of cranial 
characters.) 



22 MR. EDAVARD DEGEN ON THE 

The Postfrontals, which are joined laterally to the single parietal 
bone by means of ordinary sutures, and which extend the whole 
length of the parietal as far backwards as the supratemporal 
bones, complete the cranial roof. They form a single plate in the 
present species, with every trace of their former sutures com- 
pletely obliterated through fusion of the two component parts 
of which they consist in a great number of Lizards. This is a 
condition found to exist also in Laceria simonyi, L. atlantica, 
L. ocellata, L. viridis, L. galloti, L. dugesii, and L. Icevis (in 
the latter sj)ecies the squamosal also fuses with the postfrontal 
plate). In Lacerta jacksoni these bones are on the same hori- 
zontal plane with the parietal to their outermost margins, which 
in so many of the Lizards of the " imoralis " group are strongly 
convex and appreciably bent downward towards the temporal 
foramen., 

A similar feature, which imparts such a quadrangular 
aspect to this portion of the skull, is observable in the case 
of Lacerta simonyi* , L. atlantica^ L. ocellata, L. viridis, L. galloti, 
L. dugesii, L. Icevis, Algiroides nigropunctatus ; and among 
L. viuralis forms in the varieties nigriventris and serpa. 

Frontal. — Originally paired (in very young specimens) it is 
single in the skull before me as well a,s in the majority of the 
members of this genus when adult. Its interorbital width, at 
the narrowest part of its anteiior portion, measvires exactly one- 
half of its posterior and widest portion, which is to be found in a 
line drawn between supraocularia 4, the latter coinciding approxi- 
mately with the fronto-paT-ietal sutui-e. In proportion, therefore, 
this interorbital width is greater than in most of the muralis-\\kQ 
forms, except for the skull of a male specimen of the typical form 
from Vienna, which is also greatly depressed. In the length of 
the skull this dimension is contained six times ; whereas in the 
majority of skulls belonging to the muralis group it varies 
between seven and eight times. It fm^ther equals the width of 
supraocularia 2 and 3 ; also that of the space between the innei* 
dentary ridges (laminae horizontales) of the maxillary bones 
(measured across the apertura narium interna), and is of the 
same length exactly as the columella cranii (epipterygoid, Parker). 
Equally distinguished in respect to this broad interorbital 
diameter are — apart, again, from all the previously enumerated 
Atlantic Island forms and others named — the skulls of L. agilis 
and Icevis, and of the more typical viuralis forms those of 
L. cJdorogaster, L. saxicola, and L. derjugini f. Further, in 
Algiroides nigrop>'anctatus this region is also of a broader type ; 
but according to Siebenrock J it differs in the persistence of the 

* Cf. " Das Skelet der Lacerta simom/i Steind. und dev Lacerfcenfamilie iiber- 
liaupt," vcn Friedrieli Siebenrock, Sitzuiigab. Kais. Akad. der Wissenschaften iu 
Wien, vol. ciii. part i., April 1894, Taf. ii. figs. 8 & 11. 

t Cfr. L. von Mehely, " Materialieii zu einer Sj'stematik und Phylogerie der 
Mnralis-aluiliclien Lacevten," Ann. Mas. Nat. Hungarici, Band vii. 1909, tab. xix. 
fio-. 9 & tab. xxii. figs. 3 & 6. 
■ji.c^p. 23. 



CHAXIAL CHARACTERS OF LACERTA JACKSOXI. 



23 



two original parts composing the single frontal in the adult of 
this genus also *. 

Text-fiiT. 6. 







so. 2.. 



SO. 3 



A. LumiwA s\\\->rR.c\\\a.Yh of Lacerfajacksoni, $. Right side. 

B. Supraorbital bone of jC.,/acl-so»;, $. Right side, ventral aspect. 

C. Supraorbital bone of i.,;arA;.50»<', ?. Left side, ventral aspect. 

T)-I. Showinar configuration of nasal process of prsemaxillarj' and outer nasal 
apertures of — (D) L.jacksoni, S \ (K) L. Jacksoni, $ ; (F) X. miiralis, f. typica, 
S , from Vienna; (G) L. Diurcdis, f. typica, (? , from Turin; (H) L. vivijwra, S i 
(I) L. ac/ilis, 3^. X6i. 

pr.f., prasfi'ontale : so., scuta supraocularia (1-4); sc, scutum supraciliare ; 
s.r., scutum retrociliare; s.or., supraorbital. 

rrfemaxillary. — Its hackwardly dii-ected nasal process (text- 
fig. 6, D) is rather short and exceptionally broad at its base, 
where it is widely expanded on both sides, close behind the 



* The skull of the mak specimen of this sperirs in the British Museum Cnllections 
does not bear out this statomcnl, the said suture being: wholly ohlileraUd hy 
incrustation. 



24 MR. EDWARD DEGEN ON THE 

laterally situated nerve-canals which at this point jjerforate the 
bone. Other examples of a very broad nasal process of the 
prsemaxillary are to be found in Lacerta ocellata, L. viridis, 
L. agilis (text-fig. 6, I) ; and reference to Dr. Siebenrock's 
text and figures for L. simonyi * also shows the latter to belong 
distinctly to the broad type, and consequently not — as he points 
out, when referring to this feature^ — being distinguished by 
a " long and slender " type of processus nasalis the same as in 
Eremias arguta, with which he likens L. simonyi as regards this 
charactei'. 

In Lacerta dugesii I find the shape of the nasal process occupies 
a position intermediate between L. jacksoni, the species here dealt 
with, and those possessing a nasal process of distinctly slender 
and elongate shape. Most of the true imtralis forms belong to 
the latter (text-fig. 6, F & G). Lacerta echinata, L. vivipara 
(text-fig. 6, H), and L. loevis must also be considered intermediate 
forms in this respect. Allowance for individual variation, how- 
ever, must be made, as the examination of three skulls of the 
latter species, all males, clearly demonstrates. 

The very low angle at which this nasal process rises at its 
issue from the maxillary portion of the bone is merel}' due to 
the strongly depressed type of the skull itself, as, in fact, in 
a,ny of the typically platycephalous forms, e. g. L. mosorensis or 
L. oxycephala, and differing from the more or less convex types 
of skulls in which this process is more highly arched. 

JVasal hones. — Owing to the profuse incrustation of the upper 
surface of the skull, their contours in this specimen are very 
ill-defined. All that can be said on this head is that they 
are oblong, of a scaly nature, and that they widely diverge 
behind to receive between them the anterior and pointed pro- 
cessus nasalis of the frontal bone. Their length approximately 
equals their combined width at the widest part between the 
prasfrontal bones. 

The Prefrontals also appear very irregular in their upper out- 
lines and in their relations to the maxillaries, the frontal, and the 
nasals. The spinous posterior process, which forms the anterior 
border of the margosupraorbitalis, seemingly does not reach much 
farther backward here than the suture between the first and the 
second supraorbitals. 

The Lacrymal bones are clearly defined, exceedingly thin, 
narrow, and blade-like in shape. In their situation at the 
anterior angle of the orbital cavity, they foi^m, together with the 
incisura lacrymalis of the prpefrontals, a well-developed foramen 
lacryviale. 

Jugulars. — Their zygomatic processes are represented by a 
rather short, but acutely pointed spine. Their temporal processes 
are somewhat straighter than is the case in Lizards having 
deeper skulls, in all of which they usually are of a more sigmoid 
form. 

* i.e. p. 32, Taf. ii. tig-. 8. 



CRANIAL CHARACTKliS OF LACERTA JACKSOM. Z^) 

Occipital Segmexx. — Sapraoccipital. Its epiotic portion is 
nearly horizontal and only slightly inclines towards the posterior 
free edge, which forms the vipper margin of the foramen magnum. 
The processus ascendens (processus spinosus, Clason) rises rather 
abruptly and almost vertically from the anterior portion of the 
supraoccipital, and constitutes the exclusive su^'iport of the 
parietal roof. 

The processus ascendens is not flanked on either side by a.uy 
apophysial eminence of the supraoccipital, which so frequently 
acquires the same height as does the spinous process itself, in 
order to increase the connecting-ai'ea with the parietal, such as is 
the case in Lacerta oxi/cephala, L.saxicola, L.cleJiUpjyi*^ L. chloro- 
gaster {boettgeri Meh. ), and L. iiiosorensis^ among some of the flat- 
headed species ; or as in Lacerta muralis forma typica, the 
varieties tiliguerta, campestris, nigriventris, &c., as well as Lacerta 
vivipara, among the more highly and convexly-roofed forms 
where it also occurs. 

A similar pillar arrangement as pointed out for Lacerta jack- 
soni is present also in L. simonyi, where it is carried out to 
even greater perfection owing to the increased elevation of the 
parietal roof f ; in L. galloti, L. vhHclis, L. diu/esii, and Z. echinata, 
proportionate, of course, to their respective dimensions of height. 
This arrangement is of rare occurrence in a strongly depressed, 
practically platycephalous Lizard, and equally so in the case of 
the more pronounced pyi'amidocephalous forms, amongst which 
I find it occurs (in two specimens) in the varieties serpa and 
lilfordi onl3^ 

The Basioccipital X and Basisphenoid § in no essential deviate 
in their configuration from the outlines usual in the ^enus for 
this region. 

The ParaspJienoid is feebly ossified at its base, whereas its 
centre remains cartilaginous. 

The Orhitosphenoid bones have a broadish appearance on 
a,ccount of their reduced size, the latter being due to the depressed 
membranous portion of the ci'anium. 

The 6'qicamosals, usually slender and delicately shaped, which 
in so many of the smaller forms of iMcerta are only loosely con- 
nected by ligamentous tissue with the outer postfrontals, are here 
strongly developed and anteriorly quite fused to the latter. 
Their posterior ends, which articidate with the quadrates, are 
broadly expanded and, together with the parietal processes and 
the much reduced supratempoials which wedge themselves 

* Cfr. Mehelv, " Materialien etc.," Ann. Mus. Nat. Hung., Bd. vii. 1909, tab. xvi. 
fig. 6, tab. xix. figs. 3 & 10, tab. xxii. fig. 6. 

t Cfr. Siebeurock, 1. c. Taf. ii. fig. 9. 

X Posteriorly and slightly in front of the foramen magnnm there is a small 
tubercular process in the male specimen, which is absent in the female. 

§ The foramen by which a branch of the jugular vein leaves the skull, and which 
Siebenrock {I.e. p. 11) regards as the result of synostosis of a special process with 
the anterior angle above it in the case of the Canary Island Lizards, luit as alisent 
in the European forms as well as in L. dui/csii, is represented in L.jacksoni by 
ligament only. 



26 MR. EDWARD DEGEX OX THE 

between these two latero-posterior angles of the skull, unite 
with the processus parotici in such a way that the foramen supi'a- 
teniporale, conspicuous for its relative large size even in the 
adults of the pyramiclocephalous forms like Lacerta muralis 
varieties Jiumana, serpa, canvpestris, piti/usensis, and others, 
is here nearly closed up as much, for instance, as in L. Icevis. 

The Prces2)henoid, which in so)ne Lacertce is a slender ossifica- 
tioii of the ethmoidal region in the membranous septihm orhitale, 
is liere reduced to a small cartilaginous band in which calcifica- 
tions in the shape of granules are imbedded. 

The Colitinella cranii, for a skull of the inferior dimensions 
of the present one, ai'e exceptionally thick and strongly bent 
laterally — particularly so at their parietal ends, where they are 
rh close contact with the ala sphevoidea. Thus they differ 
somewhat from the more delicate, either linear or slightly sig- 
moid thin structures frequently met with in the majority of other 
Wall-Lizards ; but this condition is the rule in the very young, 
as already pointed out by Siebenrock *. 

The Quadrata and Ossa transversa both answer to the general 
form of description for a\\ these Lizards. 

The Vomer plates are closely approximated to each other 
medially along their entire length, but intimately united only 
at their anterior ends. The lacuna pterygo-vomerina does not 
extend beyond the most posterior portion of the palate-bones, 
which latter also unite with each other along their greater length, 
producing the crista vomerina which is continued on to the 
palatines. 

Mandible. — This consists of five elements only, a condition stated 
by Leydig to be typical of the Lacertoi, and further confirmed also 
by Siebenrock t in the fully adult Lacerta atlantka, L. muralis 
var. ruelissellensis, L, ccerulea^ L. oxycepliala^ and L. mosorensisy 
a condition brought about by the fusion of the supra-angulare 
with the articulare ; whereas in L. simowji, ocellata, galloti, 
and viridist, according to Siebenrock, these tw^o bones remaiix 
separated throughout life. 

The number of teeth in each ramus is 23. 

Description of the Skull of the Female. 

Dimensions. — Length. 15 mm. ; width 9 mm. ; height 4 mm. 

Its width is therefore contained 1-6 times in its length, and its 
height 3 '3 times. 

General Configuration. — Strongly depressed in the same pro- 
portions as in the male. 

In textui'e the bones are luther delicate, such as is character- 
istically the case in all platycephalous species of Lizards so 

* L. c. p. 52. t L. c. p. 53. 

J In the skull of a specimen of this species from Ho/.en, Tyi'ol, tliesp two bones, 
though fused slightly, show their original longitudinal sutures in their anterior 
portions. 



CRANIAL CHARACTERS OF LACERTA JACKSONl. 



27 



typically represented by Lacerta mosorensis, oxycephala, &c. 
Except for the bones composing the occipital segment, and the 
iisually more massive preemaxillary and maxillaries, all the others 
assume a more or less foliated form. 

Facial portion. The snout appears much shorter and broader 
than in the larger male, but is slightly more pointed in front. 



Text-fig. 7. 







Skull of female. 
Upper, lower, lateral and posterior aspects. X 3j. 

Cranium {Fronto-parietal region). — Slightly broader than long 
in proportion, owing to the incomplete development of its posterior 
margin, which is crescentic in shape as, in all young examples. 
Its median edge therefore barely reaches the anterior margin of 
the supraoccipital. The upper surface, quite horizontal in its main 
configuration, is longitudinally slightly undulated by alternately 
being concave in its fronto-nasal and fronto-parietal parts re- 
spectively, and convex in the intervening fi-ontal and pai'ietal 
areas. 

Dermal incrustatio7i. Very sparse and transparent. The sulci 
of the blood-vessels are sharply defined for their former rami- 
fications. 

Lamina supraciliaris. Completely ossified also, as in the male, 
and not bearing a trace of a membranous fontanelle. 



28 MR. EDWARD DEGEX ON THE 

Suprao7'hitals (text-fig. 6, C, p. 23). More regularly trinngulnr 
than in the male, also slightly more massive. Their thickened and 
strongly convex lateral margins, scarcely exposed in the male, are 
here so to a greater extent on both sides {ascertained before 
accidental displacement of their dermal plates on one side in the 
later manipulation of the skull), having been covei'ed partially 
only by the supraciliary. 

Pterygoids. Diverging slightly more than in the male speci- 
men. Teeth absent. 

Postfrontals. Forming single plates also. Faint traces of an 
earlier existing sntnre, as recorded to exist in the very young 
stages of Lacerta dvgesii, ocellaia^ and muralis by Siebenrock*, 
are discei'nible anteriorly only. In their posterior portions 
(postorbital margins) their fusion is complete. 

A minutely developed Retrociliary present. 

Dermal ossifications of Temporal region. I have been unable 
to find a,ny. 

Dentition. Number of teeth in nrjemaxillaxy 9, in each maxillary 
15-16. 

Nasal apertures (text-fig. 6, E). These are slightly narrower 
than in the male, and therefore longer in proportion. 

Parietal Region. — Except for its morphogenetically inferior 
development of the posterior margin, which is a common feature 
of all the young, this does not deviate in any essential from the 
usual conditions. Its relation to the supraoccipital alone requires 
some explanation (see p. 29), which latter bone is, owing to the 
development stage, wholly exposed and not yet bridged over as 
it is m the male. 

The Foramen parietale is absent, as in the male. 

Frontal. — The median suture is discernible in its greater 
length, which is quite in accordance with the remarks alieady 
made on this subject. The width of the frontal at its narrowest 
point (between the margines orbitaJes) is equal to that of supra- 
oculai-y 3. The frontal is actually and I'elatively narrower than 
that of the male, its width being slightly less than half the 
length, and a seventh of the total length of the skull, whilst in 
the male the similar measurement shows that the frontal is 
relatively larger. The width of tlie female frontal is three- 
fourths of the length of the columella cranii, and in this measure- 
ment, as well as when it is compared with the width between 
the internal laminse horizontales (taken diagonally across tlie 
choange), the relative and actual inferiority of size as compared 
with the male is shown. 

Praemaxillary (text-fig. 6, E). — The nasal process of this bone 
is on the whole longer and narrower than in the male. It alFo 
terminates posteriori}^ in a much more finely drawn-out point. 
It is quite devoid of any calcareous incrustation. Anteriorly it 
rises from the maxillary portion of the bone at a similar acute 

* i. c. p. 75. 



CRANIAL CHARACTERS OF LACERTA JACKSONI. 29 

atigle to its hoi-izontal plane as in the male, but is less expanded 
at its base, immediately beliind the constriction between the 
nerve-holes. 

The Nasal bones, which, owing to extensive incrustation, in 
the male specimen were described as ill-defined, are here quite 
distinct in their entirety. Anteriorly their proximal processes, 
between which the nasal process of the praemaxillary is wedged, 
extend to the middle of the latter. Their lateral margins, shorter 
than those forming the median suture and moreover less decidedly 
linear, are parallel to each othei'. Their posterior edges, scale- 
like in shape, and imbricating in a similar manner on the fi-ontals, 
are characteristic in having from 2 to 3 pointed lobes, of which 
the median ones are the longest, so as to become a fork into the 
prongs of which the median process of the frontal penetrates. 

The Prcefrontals also ara more clearly defined in the female. 
They show their posterior frontal processes, which constitute the 
margo orbitalis anteriorly, to be spurs which extend nearly as far 
back as the suture between supraocularies 2 and ?>. 

The Jugulars do not dift'er from those of the male. 

Occipital Segment. — The tSujn'aoccipital is united along its 
entire upper margin with the posterior edge of the parietal bone. 
The processus ascendens, so tolerably well developed and forming 
the sole support of the parietal in the male, is quite rudimentary 
here and reduced to a tubercle only. It is lower even than are the 
a.pophysial elevations of the epiotic portions of the bone, and, 
together with the epiphysial cartilaginous spine and a small 
portion of the membi'anous cranium, is laid bare and quite un- 
protected by the paiietal. The sutures between the supraoccipital 
and the pleuroccipitals are perfectly distinct. The foramina 
retrofrontalia, in consequence of this extended transverse con- 
nection between the parietal and the supraoccipital, are more 
markedly reduced than in the male. 

The Basioccipitcd and Basis2)henoid are not fused as in the male, 
the suture between them showing distinctly (text-fig. 7, p. 27). 

The Squamosals are flatter and less curved downwards than in 
the male. 

The Vomer ^^Icties are more loosely united than in the male ; but 
the Palatines, though more distinctly united, have their sagittal 
ridges imperfectly developed as compared with those of the 
male. The stolci palatince are also shallower in this skull. 

Mandible. — This consists of the six elements, of which each 
ramus is composed in all immature Lizards as pointed out 
by Siebenrock, the supra-angulare and articulare being not yet 
united by fusion. 



On the Absence of the Foravien parietale. 

After first ascribing the absence of the orifice for the pineal 
eye in the skull of the male to purely pathogenic causes, or to 



30 MR. EDWARD DEGEX OX THE 

a possibly individual abej-ration, both of these assumptions were 
dispelled on the discovery of this peculiarity in the female also. 

An examination of the remaining five specimens preserved 
in alcohol, as well as of the type specimen referred to (antea, 
p. 20), and of a second female moi^e recently described * (all 
in the Collection of the British Museum), readily confirmed its 
specific importance, since not one of them bore the least trace of 
the presence of this morphological feature. 

Besides the simultaneous observation of its absence in Zono- 
saurus niadagascariensis, belonging to the neighbouring family 
Gerrhosauridse, a more methodical search among the rich 
material of the Biitish Museum Collections revealed the following 
genera and species belonging to the family of Lacertidfe as being 
devoid also of the foramen parietale : — 

Poromera fordii, Benito River, Spanish Guinea. 

lAtcerta echinata. Kribi River, Cameroon. 

Algiroides africamis. Uganda. 

Nucras delalandii. Natal. 

Latasiia hm-deggeri. Somaliland. 

,, neumanni. Arabia. 

,, degeni. Somaliland. 

,, spinalis. „ 

,, phillipsii. „ 

GastrophoUs vittata. Zanzibar. 

Holaspis guentheri. Benito R., Spanish Guinea. 

As the above list shows, cases of absence of the parietal 
foramen occur in seveial monotypic genera, also in the specifically 
numerous genus Latastia in no less than five species out of ten. 
Yet it is rather remarkable to find such a character restricted to 
two species only in a large genus such as Lacerta. 

But more remarkable, perhaps, is the fact that all the forms 
thus deprived of the foramen belong to the African Continent, 
aiid that in the not strictly African genera it is restiicted to 
A f rican representatives. 

Some of the remaining genera of this family, even of purely 
African habitat, do not contribute any specific forms which exhibit 
this negative character. It is curious to note, moreover, that 
among the upwards of thirty known species of the genus Eremias, 
about twenty of which are found in Africa, no exception to the 
presence of the parietal foramen could be found. 



The sum total of cranial characters, as shown piincipally in the 
immature skull of the female, points to Lacerta jacksoni being 

* G. A. Bonlenger, "Ruwenzori Expedition Reports," Trans. Zool. Soc. vol, xix. 
part iii., Dec. 1909, p. 242. 



CRANIAL CHARACTERS OF LACERTA JACKSONI. 31 

clasely allied to L. muralis, forma typica. It agrees with the 
latter as well as with L. mosoreiisis in the strongly depressed and 
more broadlj'^ expanded skull-structure in general. Also in the 
widei- interorbital region, likewise a marked feature in Lacerta 
vivipara. Also in the mode in which the connection between the 
posterior portion of the parietal and that of the supraoccipital 
presents itself ; in the absence of teeth on the pterygoid bones, 
and in the degree of divergence of these latter, as well as in the 
configuration of the basioccipital and the basisphenoid. Fiuther, 
in the shortened muzzle with broadened base, and in the 
anteriorly bi-oadened nasal process of the pnemaxillary bone 
(text-fig. 6, D), well indicated in the typical form of L. muralis 
from Turin (text-fig. 6,G), but assuming its widest proportions in 
Lacerta agilis (text-fig. 6, I). 

The total absence of supraocular fontanelles, notwithstanding 
the typically platycephalous structure, together with the onto- 
genetically early complete fusion to one composite plate of the 
postfrontals, would justify our referring this type of skull to the 
group of Lizards to Avhich the term of Neolacei'tte * has recently 
been given. 

However, considering the views held and recently enunciated 
by writers on the subject of the phylogenetic relation of some 
of these characters, it is luther surprising to find undivided 
postfrontals also in species belonging to entirely dift'erent genera, 
such as Eremias, as Prof. v. Mehely t regards that genus as one 
of the prototypes of ai'chfeolacertic forms of this family because 
of the character of its lepidosis. 

I might add that in Acant/iodactylns hoskianusX, a lizard to 
which Siebenrock § ascribes a divided condition of the post- 
frontals, I find them to be completely fused into one plate in the 
skull of a female in the British Museum Collections. 

A perfectly obliterated suture between the two postfrontals 
I also find in a female specimen of Scapteira knoxii, another 
decidedly platycephalous species. 

Thus, beyond some more or less doubtful points in regard to a 
satisfactory solution concerning the question of phylogeny, skulls 
of females do not seem to afford trustworthy indications from 
which to draw conclusions as to affinities. It is to that of 
the male, so far as an examination of the present limited 
material is concerned, that one must turn for better results. 

As pointed out in the detailed part of the description, the 
tabular surface (inclusive of the postfiontals) is a very conspicuous 
feature of this male skull when taken in conjunction with the 
quadrangular and posteriorly well projecting parietal region. A 
near approach in this respect is to be found in a male specimen 
of Lacerta muralis of the variety lilfordi, a form restricted to 

* L. V. Mehelj-, Materialieii, 1. c. 

t li- X ^^"^ "ot stated by Siebenrnck. 

§ Cf. " Das Skelct der Lacerta simoiiyi" Sitzb. Akad. Wis!<. Wieii, 1894, p. 37. 



32 MR. EDWARD DEGEX OX THE 

the Balearic Islands. Though unquestionably more pyramido- 
cephalous in regard to its supei'structure, it nevertheless holds a 
position somewhat intermediate between L. jacksoni and some 
of the more pronounced pyramidocephalous forms in this group 
in regard to this particular feature. 

The variety lilfordi also shows the same exclusive pillar arrange- 
ment of the processus ascendens for sole supjDort of the parietal 
roof, which is so prominent a feature also in all the larger foiins 
of Lizards from the Atlantic Islands such as L, sivionyi, gaUoti, 
atlanticcij and also L. dugesii * (except in regard, to a modification 
in the basal portions of the parietal process), all differing by having 
pterygoid teeth, and all of which, except the latter, have strongly 
ossiiied temporal regions. These dermal ossifications in Lacerta 
jacksoni, as may have been seen, are present in a rudimentary 
state in two of the supratemporal scutes. Specimens of Lacerta 
viridis, agilis, and kevis may be quoted as further examples of 
characteristic forms in which the postfrontals are single plates, 
with perfectly analogous conditions in the relations between 
parietal and supraoccipital. 

Also Lacerta ecMnata, from West Africa (except for its more 
elongate skull), in which the processus ascendens, typically pyra- 
midal, does not enter into direct contact with the parietal, but is 
actually separated from it by confluence of the foraminpe retro- 
frontalia. 

But it is the shape of the nasal process of the pr?emaxillary 
perhaps, as previously remarked on in the male, which may 
also provide a likely indication of closer affinity with some of 
the Lizards of the Canary Islands and the European species of 
Lacerta, viz. ocellata, viridis, and agilis — a character in which the 
Balearic variety lilfordi does not share, being distinguished by a 
narrower and very slender nasal process. 

So far as the topography of the skull reveals phylogenetic 
affinity, it would appear that Lacerta jacksoni came from a 
Palfearctic stock, but also exhibits special relationships betw^een 
the West African and Atlantic Island forms. 



Remarks on Characters of Lepidosis in Lacerta jacksoni. 

Dimensions. — In regard to size, five specimens, all of them 
males, are larger than the type of the species ; the two biggest 
measuring 86 mm. from snout to vent, and consequently of 
about the same length as is the largest of the males of Dr. 
Peracca's series (consisting of seven specimens also), the length 
of which is given as 85"5 mm. 

The male specimen, from which the skull was prepared for the 

* Siebenrock {vide I. c. p. 36) includes this species amongst those having divided 
postfrontals. The specimens at my disposal, from the British Museum Collections, 
show them distinctly to be single plates, with no indication of former sutures. 



CRANIAL C'HARAf'TKRS OF LACKRTA JACK.SOXI. 33 

foregoing descriptioii, laeaisured 70 uun. : equal to tlie length of 
the type .specimen. It is therefore only slightly longer than the 
>skull of the female which was sacrificed for the sfinie purpose, the 
length of the latter from snout to vont being 67 mm. Though 
diffej-ing hy 3 mm. only, the sexual discrepancy in the size of the 
heads is a striking one : 19 nun. for the uiale against 14 mm. for 
the female. 

Analogous specimens for sfze in Dr. Peracca's series show 
corresponding difterences, viz. 20 mm. for the head of a male 
and 15-5 mm, for a femnle. 

Ventird Plates. — The number of tlie ventral plnte series varies 
in the Mount Elgon specimens, three of these having 6 longi- 
tudinal rows only and four of them having 8, which is the same 
as in the type of the species. 

For the Turin Museum specimens their number is uniformly 
stated to be 6 ; but to judge from a footnote in connection with 
this point, the author was fully cognisant of the presence of 
supplementary series* for some of his specimens, but hesitated to 
accord them serial rank on account of the inferior size of the scutes 
which form the outermost rows. 

In several of the Mount Elgon specimens these latter are 
sufficiently well developed to be considered as a series of venti^als. 

TrauvSversely, the minimum of the plates in the Elgon speci- 
mens is 23 in an adult male, and the maximum 26 respectively 
for the male and the female from which the skulls ha.ve been 
prepared. 

For the Turin Museum series this minimum of 23 occurs in 
two males, one quite young and the other larger, whereas for 
the largest and fully adult the number is 24, but the highest 
figures of 27 and 28 are reached by two females of the same lot ; 
the first number being identical with that of the Ruwenzori female 
in the British Museum Collections, whereas the female specimen 
from Mount Elgon has only 24 rows of these scutes. 

It must be borne in mind that female lizards, having the body 
more elongate in proportion than the males, usually possess a 
higher number of transverse series of ventral plates. 

/Scales across the middle of the Body. — These range between 
38 and 43, giving an average of about 40, the same as recorded 
for the type specimen, j^rovided we except the male from which 
the skull was prepared and which showed the abnormally high 
number of 49. llie Ruwenzori female has 37, a number 
close to the average of 36 as represented in the Turin Museum 
series, where they vary between 34 and 38, the latter figure 
being that also of Lacerta vauereselli, as well a.s constituting 
the minimum of the British Museum Elgon series of Lacerta 
jacksoiii. 

Supraciliary granules. — These vary in our specimens, both 
individually and for the two sides, between 3-3 and 6-5. They 

* Cfr. Dot-.t. M. G. Peraccn.. l\ Ruwcnzon, Relazioni SoientificlK^, vol. i. Zoologia- 
liotanica. : Rettilicd Aiuiibii, Laoortidae-, pp. 16(5 i lt)7. 

Proc. Zool. So<:.— 1 91 ] . Xo. 111, 3 



34 MR. EDWAHL) DEGEN ON THE 

are confined principally to the posterior portion of the supraocular 
region. In the Ruwenzori female they are i-educed to 3 on the left 
and to 2 on the right side. Di-. Peracca describes these as " very 
ii'ragular and asynametrical for the two sides, except in a female in 
which the seiies is a complete one on the left, but totally absent 
on the right side." Accoixling to Tornier, who in his description 
omits to quote their number, the supi'aciliary series of granules is 
incomplete also in Lacerta vauereselli ; but, as he says, confined to 
the posterior portion, agreeing therefore in this point also with 
L. jacksoui. 

Galar Scales. — The number of scales between the third chin- 
plates and the collar-plates varies between 24 and 26 in the Elgon 
specimens except in one, in which there are 28. This is slightly 
in excess of the Duke of the Abruzzi's Expedition specimens, in 
which they are stated to be 22. In the type of the species 
their number is 25. 

Femoral Pores. — Their number is fairly constant, Avith variations 
between 16 and 18 in the Elgon series. In one of the two females 
of the Turin Museum series there is a red uction to 15-14, otherwise 
17-18 being the normal number. No case of supernumerary pores 
occurs in any of the British Museum specimens similar to the one 
female quoted in this respect by Dr. Peracca,, 

Upper Labials {^anterior to the siobocular). — Except for the head 
of the male, made into a skull, which on its left side showed these 
to be 5, their normal number throughout the remainder is 4. 
A male of Dr. Peracca's series shows a similar variation. 

Postocular and Temporal Region. — In one of the specimens the 
last postocular scute is in contact with the first supra-temporal, 
similar to what is found in Lacerta inosorensis as figured by 
Mehely *. In all others it is the upper postocular and the parietal 
scutes which meet, as in Lacerta horvathi Meh. or~Z. miiralis 
typica t. 

The 3 to 4 supratemporals are exceedingly small and nari-ow, 
the first and longest being partially supported proximally by 
the lateral margin of its underlying postfrontal bone, and 
therefore visible practically in its greatest extent when the head 
is viewed from above. 

The number of scutes for the temporal region varies greatly. 
Those, for instance, forming the second row between the large sub- 
ocular and the first supratemporal are : 3 for the left, a.nd 5 
for the right side in the male, the skull of which served for 
description. 

Similar variation is to be found amongst the scutes in regard 
to their sculpture, which shows every gradation, from the per- 
fectly smooth through the granular stage, as pointed out in Mr. 
Boulenger's description of the typical specimen J, to the faintly 
keeled or tuberculai' scutes to be met with in some of the Elgon 

* Cfr. Ann. Mus. Xat. Hung. ii. 1904, p. 366, fig. 2, A, "Eine iieue Lacerta aus 
Ilngarn." 

tJft. %. 3, B&C. 

% Cf. G. A. Boulciiger, Proc, Zool, Soc. 1899, p. 96, 



CRANIAL CHARACTEUS OF I.ACERTA JACKSONI. 35 

specimens, a condition said by Tornier to exist also in his Lacerta 
vanereselli. 

The suture between the first supraciliaiy and the second is 
slightly oblique. 

The frontal shield, which also varies slightly in length, and 
which is especially short in some specimens, is on the whole shorter 
and broader than would appear from the figure on the plate 
accompanying the description of the type of the species. It 
was particularly bi'oad in the male specimen which supplied the 
skull for description. 

Colour-pattern and markings are quite uniform for the Elgon 
specimens, and identical with the type of the species. In their 
nearly fresh state of preservation their ventral sides were of a 
vivid yellow, which, since their preservation in alcohol, has changed 
into the whitish blue-grey described by Tornier in his Lacerta 
vauereselli, or the pale blue ascribed to the Ruwenzori female. 



Reraarhs on some of the sicpjwsed Specl/tc Characters of 
Lacerta vauereselli Tornier, 

On examination of specimens in a comparatively large series, 
as may have been seen from the foregoing, it appears that some 
of them are subject to individual variations. 

This is particularly noticeable in the scaling of the temporal 
region, which was made one of the principal features by which 
to distinguish Lacerta vauereselli from L. jacksoni. 

In the majority of specimens of the la,tter, these scales cor- 
respond with Dr. Tornier's description of them *, and a special 
note on this subject made on the male specimen — previous to re- 
sorting to the excision of the skull — was to the efi'ect that these 
scales were considerably larger than those of the upper dorsals, 
being oblong, irregular in size, hexagonal to pentagonal in 
shape, feebly though distinctly keeled, and decidedly double the 
size of those situated immediately behind the auditory opening — 
all of these being the chnracteristics ascribed by Dr. Tornier to 
his Lacerta vauereselli^ and which are also referred to by Mr. 
Boulengei' t at the end of his description of the Ruwenzori female. 

According to Dr. Tornier the collar is serrated. This, again, is 
a point not always easy of recognition, for some specimens show 
various degrees of unevenness in this respect. 

The same rema.rk applies to the number of plates forming the 
colla.r-edge, which are stated by Toraier to be 11. In the type of 
the species they are stated to be 10, which is the number also in one 
of the Elgon specimens ; in the remainder of which, however, 
they range downward to from 9 to 7 only, thus showing con- 
siderable fluctuation. Pei-acca omits to quote their number. 

* Cfr. Zool. Air/.oiger, Bd. xxv., Oct. 1902. p. 702. 

t " Ruwenzori Exped. Reports," Traus. Zool. Soc. vol. xix. part iii., Dec. 1009, 
p. 242. 

3* 



36 UN THE C'KANTAL CHAKACTERS OF LAC'EliTA JACKSO.VI. 

iS^or can I find any differences in the istyle of scaling of tlie 
upper dorsals between Dr. Torniei^'s diagnosis and that given 
for the type of the species by Mr. Boulenger, which scales, according 
to the latter, and further verified by myself, are stated to be 
" rhomboidal, keeled, juxtaposed, or subimbricate," eveiy one of 
these conditions occurring in the Elgon specimens as well as in 
Lacerta vauereselli. 

The number of scales aci-oss the body, viz. 38 in the latter, also 
is in accord with the mean of the Elgon series of specimens ; and, 
as regards the femoral pores, this has been finally disposed of, 
both by what has been made known [antea, p. 34), as well as by 
a remark made by Mr. Boulengei' in connection witli this point 
in his description of the Ruwenzori female. 

As already pointed out in the diagnosis of the skull of Laceria 
jacksoni, there are no teeth on the pterygoids any more than 
in L. vauereselli. No difference appears to exist in I'egard to 
the supratemporals as described for this species by Dr. Tornier 
fi'om their condition in the Elgon specimens, in all of which a small 
tympanic scute is plainly visible. 

Neither is a distinct gular fold recognizable in all specimens, at 
least in the stricter sense, as it is often apparently absent in im- 
mature individuals, such as seems to be the case with Dr. Tornitr's 
type, judging b}' the dimensions given by him — an assumption 
which is further corroborated by the fact that its sex has not been 
mentioned. 

The length of the posterior extremity also varies gieatly 
individually — in some Elgon examples it scarcely attains the 
axillaiy^ pit, whilst in others it reaches well beyond the collar- 
edge. 



The geographical range of this species, so far as the increased, 
but at piesent still scanty, material on I'ecoid permits one of 
judging (17 specimens, including Dr. Tornier s L. vauereselli), 
appears to be a strictly equatorial one. 

Its habitat, moreover, seems to be generally restricted to the 
mountainous region of that part of the Central African plateau, 
or immediately west of it, which culminates in its highest peaks, 
an area generally acquiring considerable altitude. 

Its ultraplatycephalous shape, concurrently with its sombie- 
coloured garb, at once seems to denote a form occupying high 
ground, differing from those of the plains, with generally more 
massive skulls and more vividly contrasting colour- patterns, 
in the same manner as, for instance, Lacerta mosorensis differs 
from Lacerta muralis var. cav}2^esiris or serpa, and L^acerta muralis 
var. monticolO' from the var. hocagii. 



00 



:/ 




u 

< 

W 
o 

O 

w 

H 
h1 



x 



ox THE PKLOI'OXMCSTAN LIZARD. 37 



3. On the Peloponnesiaii liizard [Lacerta peloponnesiaca 
Bibr.). By G. A. BocTLENGER, F.R.S., V.P.Z.S. 

[Received Octoher 4, 1010 : Read November 15, 1910.] 
(Plate I.* and Text figure 8.) 

Examples of a Lizard from Gi'eece, which had been confounded 
with Lacerta 7nurali!s or with L. taurica t until shown by Dr. 
J. de Bedriaga X to be fully entitled to specific rank under the 
na,me of L. peloponnesiaca Bibr. §, have been exhibited for the 
first time this summer in the Societ3'''s Reptile House. As regards 
coloration, the species is one of the most beautiful, and as the 
oidy figures of this Lizard we possess are quite inaccurate, or taken 
from specimens preserved in spirit, I requested Mr. J. Green to 
make coloured pictures from the specimens living in the Gardens ; 
at the same time I seized this opportunity of drawing uji a new 
description of this little-known Lizard, with a view to fixing its 
correct position in the genus Lacerta. 

In addition to tlie live specimens I haveha.d the following small 
series at my disposal, all except no. 3 being preserved in the 
iS^atural Plistoi-y Museum : — 

1-2. (^?. Movea. Dr. J. de Bedriaga. 

3. (J. ,, ,, (Lataste Collection). 

4-8. ^ $ ligr. L. Stj'mphalos. Norman Douglsss, Esq. 

P-10. (??. Olvmpia. Hr. Loreiiz Miiller. 

11. (?. Kalamata. 

The skull of a male fi-om L. Stymphalos bas been prepared, and 
agrees in all essential respects with that of L. taurica. 

Form and Proportions. 

As may be seen on comparing the following table of measurements 
with that given for Z. taurica\\, the general pi-oportions are the 
same as in that species, except for the rather longer limbs. In 
the male the hind limb reaches the collar or a little beyond, in the 
female it reaches the axil or the shoxilder. The head is less 
flattened than in any of the forms of L. muralis, closely resembling 
in shape that of the typical form of I/, riridis ; snout sometimes very 
obtuse, sometimes very pointed, as long as the distance between 
the eye and the ear-opening; neck as broad as the head, or a 
little hrondei'. The foot is always longer than the head. 

* For p.xpliniHlion of IJip rJHtc srr p. 10. 

+ For a description of this Li/.ard sec P. Z. S. ]!in7. p. 557. 

t Zoo]. Anz. 188:1, p. 21H, and .■\hli. Senclc. Ges. xiv. 1886, p. 31-5. 

§ In Horv de St. Vine. Expcd. Sc. Morec, iii. p. 6<i. 

I'i P. Z. S. 1007, ]). .",B:i. 



38 



MH. G. A. BOULKXGER ON 



Measurements (in millimetres). 

<? • ____J .__ 

1. 2. 3. 4. 5. 6. 7. 8. 9. l3. 

From snout to vent 80 76 76 75 71 70 77 71 70 65 

„ fore limb 32 29 29 30 28 27 28 26 25 25 

Length of head 21 20 19 19 19 19 16 16 16 16 

Wicltli of head 13 13 13 13 12 12 10 10 9 9 

Depth of head 11 10 11 10 10 10 9 9 8 8 

Fore limb 27 26 26 25 25 25 24 21 23 22 

Hindlimb 47 45 46 42 42 43 41 35 37 34 

Foot -. 25 23 24 22 22 23 21 19 22 18 

Tail (*reprodiiced) 170 95*77*100*80*140 105 97*98*125 

1, 9. Greece; 2, &. Olympia; 3, 6, 7. Morea ; 4. Kalamata; 5, 10. L. Styraphalos. 

Palatal Teeth. 

These are strongly developed, forming two to four longitvidinal 
sei'ies on each pterygoid bone. 

Scaling. 

Rosti-al shield largely entering the nostril. Nasals forming a 
short suture behind the rostral * ; frontonasal broader than long ; 



Text-fig. 8- 







Upper and side views of head of male and female (from photographs). 



frontal not longer than frontoparietals, anterior borders very 
concave, forming an acute angle, the shield often appearing 

* In one specimen the frontonasal forms a short suture with the rostral. 



THE rELOl'O.XNESlAN LIZARD. 39 

trilobate in front, in a,dnlt male specimens* ; parietals about once 
and a half as long as broad, in contact with the upper postocular ; 
occipital ver}^ variable in size, sometimes longer, sometimes 
shorter, find usually broader than the interparietal, from which 
it may be separated t by a short suture formed by the parietals. 
Supraoculars in contact with the supraciliaries, rarely one or 
two granular scales intervening between them. Postnasal single +. 
Four or five upper labials anterior to the subocular §. Rather 
large, irregular shields cover the temple, often exactly as in 
L. viridis ; tympanic shield distinct ; 3 to 5 upper temporals in 
contact with the parietal. 28 to 33 scales and granules in a 
straight line between the symphysis of the chin-shields and the 
median collar-plate ; gular fold distinct. Collar with even edge, 
formed of 10 to 12 plates. 

Scales on back juxtaposed, granular, round or suboval, feebly 
or faintly keeled, on flanks towards the ventrals larger, flat and 

Table showing number of scales, plates, and femoi^al pores. 

1. 2. 3. 4. 5. 6. 7. 8. 

<J. Greece 80 56 30 10 31 4-5 24-25 29 

„ Olynipia 76 55 28 11 30 4 23 28 

„ Morea 76 57 28 12 28 4 20-21 30 

„ Kalamata 75 60 29 10 30 5 21 26 

„ L. Stymphalos 71 63 28 10 30 4 22-24 28 

„ Morea 70 62 28 11 31 4-5 23-25 31 

?. Morea 77 59 31 11 30 4-5 22 29 

,. Olvmpia 71 53 32 11 31 4-5 23 27 

„ Greece 70 60 .33 11 31 4-5 20 30 

„ L. Stymplialos 65 62 31 11 33 4 24-25 27 

1. Length (in millimetres) from snout to vent. 2. Number of scales across middle 
of body. 3. Transverse series of ventral plates. 4. Number of plates in collar. 
5. Number of scales and granules between symphysis of cliin-sliields and median 
collar-plate. 6. Number of upper labials anterior to subocular. 7. Number of 
femoral pores. 8. Number of laniellar scales under fourth toe. 

subimbricate, smooth ; 53 to 63 scales across the body, 2 or 3 
transverse series corresponding to one ventral plate. 

"Ventral plates in 6 longitudinal and 28 to 33 transverse series. 
Anal plate moderately large, with 2 or 3 semicircles of scales 
around it. 

Scales on upper surface of leg a little smaller than dorsals, 
distinctly keeled; 26 to 31 lamellar scales under the fourth toe; 
20 to 25 femoral pores on each side. 

Caudal scales truncate behind, more or less strongly keeled, the 
whorls subequal in length. 

Coloration. 
This varies considerably according to individuals. Females and 

* In the allied L. taurica the anterior borders of the frontal vary from feebly 
concave to feebly convex, as in L. nmralis and its varieties. Peracca has attached 
undue weight to this character in his description of L. sardoa. 

t In 4 specimens out of 12. 

X Two regularly superposed postnasals in one specimen ( $ , Morea). 

§ 4 on both sides in five specimens 4 on one side and 5 on the other in six, 5 on 
both sides in one. 



40 MR. E. G. BOULE.NOEK ON 

young are beautifully sti'iped with dai"k brown or black, some 
specimens may even be descril:)ed as black above with 5 or 6 light 
longitudinal streaks. Traces of this striation may persist in some 
males, but usually disappears entirely, the back being uniform 
brownish or dull green, with small blackish spots or vermicular 
lines on the sides. The figui'es annexed to this description will 
give an idea of the gorgeous colours assumed by the adult male 
during the breeding-season, making it one of the most beautiful 
of the European Lizards. The top of the head is of a reddish 
brown, sharply contrasting with the gTeen colour of the nape, 
which gradually chaiiges to olive or brown on the postei'ior part 
of the body ; the sides of the head and body and the lowei- pa.rts 
are of a bright vermilion-oi'ange, relieved bya patch of azure-blue 
in the axillary region and a broad band of the same colour occupying 
the outer row of ventral shields and extending a, little way tip on 
the scaly part of the side. The female is of a reddish brown, 
with two broad blackish bands along each side, the outer proceeding 
from the eye, bordered above and below by a narrow whitish 
strea,k and separated by a third light strea.k which in some 
specimens is pale yellow, in others pale green ; a small round 
blue spot is present above the axil ; the hind limbs bear round 
light spots edged with blackish ; the lower parts are white or 
pale yellow, often tinged with rosy or lilac on the sides. 

Bel'itionships. 

Lacerta peloponnesiaca is most nearty related to L. tanHca. a.nd 
with it fills the gap between the massive Lizards like L. agilis 
and L. viridis and the forms that cluster round L. vitivalis. In its 
thick, convex skull and its well-developed pterygoid teeth, as well 
a,s in its temporal scutellation, it agrees with the former group, 
which I 'regard as the most gener-alized, whilst in its scaling and 
especially in its non-serrated collar it agrees very closely with the 
typical L. mic^'alis, from which some authors have held it to be 
probably derived. 

EXPLANATION OF PLATE L 
harerta pelopnnnesiara, male and female, natural size. 



4 R-einark.'' on Two Species of Fishes of the Genus Gnbius, 
from Observations made at Roscoff. By Et>ward G. 

BOULENGER *. 

f^Received Ootolier 30, If) 10: Read November 15, 191(.).] 

Prof. F. Guitel, in his well-known paper on the breeding- 
habits of Wohiiis mmu.tns'^, remarks that tl"io doscriptions of this 

* {'oniniuiii(.-Hted by (j. X. BurLUNOicn. F.R.S., V.P.Z.S. 
t Arch. Zool. Exp.'ft-t (ivn. x. 1R,Q2, p. \m. 



risHi:s OF THi: oems gouius. 41 

species bv various authoivs vaiy to a consiclei'able extent and thiit 
the fish described under this iiiuiie by Collett* does not tally 
with the said Gob}^ of Roscoff on which he based his observations, 
the number of scales along the lateral line in his specimens being 
about 45, those of the Noi-wegian fisli about 60. Further that 
Collett's formula for the fin-rays- I). 6/11-12. A. 11— diflfers 
slightly from that of the Roscoff fish, which he gives as : D. 6/8 
10. A. 8-10. 

During a stay at Roscoft' last summer I collected a number of 
specimens of the Goby describetl by Guitel luider the name of 
G. minutits and also fouiul thei'e another Goby which differed at 
a first glance from the former in its larger size, coloiation, and 
general appeai-ance, and which I found on examination under a 
lens to have more numerous scales. This is the form described 
by Messrs. Holt and Byrne t, in their paper on the British and 
Irish Gobies, as the typical Q. minuh(.s, while the fish so commonly 
found at low tide in the pools of the shallow sandy bays of 
Roscofi' is )-ega.vded by them as nu estuarine race of the same 
species, to which they refer the G. microps of KroyerJ and later 
Scandinavian authors. 

The colour of the la.tter fish is dorsally of a dirty grey, minutely 
speckled with black, laterally with lai'ge blackish blotches, which 
in the males usually expand into vei-tical bars on tlie side. That 
of the former is creamy speckled with rusty brown and with small 
blotches of the same colour laterally ; the blotches may also 
form bars, which, however, are always finer and less conspicuo\is. 
1 found the smaller form to be stouter than the larger, the depth 
of the body being usually from 5 to 6, as against 6 to 7 times in 
the total length, caudal fin excluded, and tlie scaleless area of 
the nape and back to be of greater extent. 

These two fishes I found under quite difi^erent conditions — the 
G. microps of Kroyer close inshore, the one alluded to by 
Messrs. Holt and Byrne as the typical G. mhndv.s at a locality 
north of the little island of Batz. opposite Roscoff, uncovered at 
the spring tides only. 

From the table of pai'ticula,rs of the two forms, given further 
on, it will be seen that they differ both in number of scales and 
fin-rays ; and there can be no doubt that the two fishes ai-e distinct 
and well deserve to lie I'egairled as valid species, not as I'aces 
only. 

Messrs. Holt and Byrne, liowever, are of opinion that a sufticient 
series of specimens fi'om various localities would sliow a complete 
gradation from the one ••race" to the other, and state that 
specimens from the Cuckmei-e river approach the typical form in the 
large number of scales and small scaleless area of the nape aud 

* Vii-li>n-);. Spl>li. F'nrli. rhvlstiaiiiH. \><~\. p. ICS. 

t Hpport oil llic Se;i and Inland Fislipn'p? of Ireland for llip Ychv lOOl. I'iirt ii. 
Appendix III. (1001). 

i Dhiiiii. Fiskc. i. p. Ufi ( 18;iH^lS Id). 



42 



MR. E. G. BOULKXGER ON 



back, though in form and colour they resemble the estuarine race. 
I have examined several specimens in the British Museum from 
the Ouckmere, presented by Mr. Byrne, but find that both in 
appearance as well as in the number of scales they agree with 
6r. microps. 

It seemed desirable to make sure, by reference to the original 
description, which of the two species so often confounded should 
bear the name of Gohms minutus. Tliis name was proposed by 
Pallas* for the fish {'^ Maris JBelgici" ) described in a rather 
puzzling manner by Gronoviust, who gives the nxnnber of fin- 
TSijs characteristic of G. micro'ps, while, on the other hand, the 
total length " tres uncias" (about 80 mm.) can apply only to the 
larger species. It is highly probable, however, that Gronovius 
had before him examples of both species from the Belgian coast j, 
and that he noted the number of fin-rays from the smaller fish 
and added to his description the size attained by the larger. 
Gmelin's§ diagnosis " albicans ferrugineo maculatus, ... D. 6, 11. 
A. 11 " can only apply to G. minutus of most authors. 

It is therefore satisfactory to find that no objection can be 
raised to the retention for the two species of the names used by 
the Scandinavian and other authors who have distinguished them. 

The following is a tabulation of the specimens of the two 
species from Roscoff || : — 

G. minutus. 



Length. 


Scales. 


Fin- Rai/s. 






D. A. 


55 


66 


VI. 12 11 


53 


67 


VI. 12 11 


53 


65 


VI. 12 12 


53 


64 


VI. 13 12 


50 


63 


VI. 12 12 


49 


71 


VI.— — 


49 


64 


VI. 12 11 


48 


65 


VI. 12 11 


46 


63 


VI. 12 12 


45 


62 


VI. 12 11 


44 


61 


VI. — 11 


39 


63 


VI. 12 12 



* Spicil. Zool. viii. p. 4 (1770). 

t Zoophylacium, p. 81. no. 276 (1763). 

J Specimens of both G. minuius and Gr. microps from the Belg-ian coast were 
sent to the British Museum by the late Prof. E. van Bent'den, and measurements 
of these will be found in the table at the end of this paper. The two species have 
hitherto been confounded by Belgian authors under the name of G. minutus. 

§ Syst. Nat. i. p. 1199 (1788). 

jl In these tables the length (in millimetres) is taken from the end of the snout to 
the base of the caudal fin. The scales are counted in a longitudinal series from the 
upper extremity of the gill-opening to the root of the caudal fiu, 



FISHES OF THE GKNUS GOBIUS. 

G. 1) tier dps. 



Jjcngth. 


Scales. 


Fin-jRays. 








IX A. 




45 


48 


VI. 10 10 




45 


50 


VI. 9 10 




45 


44 


VI. 9 10 




44 


52 


VI. 10 10 




43 


44 


VI. 10 9 




42 


46 


VI. 9 10 




41 


47 


VI. 9 10 




40 


49 


VI. 10 10 




40 


45 


VI. 9 9 




37 


40 


VI. 9 10 




36 


48 


VI. 1(1 10 




3i 


49 


VI. 10 10 





In order to satisfy myself as to the degree of constancy of 
these characters, I have examined a number of specimens in the 
Bi-itish Museum collection from v^arious localities, including the 
Cuckmei'e river and the Belgian coast. 

The following table may be useful for purposes of comparison : — 

G. minutus. 



Locality. 



Scales. 



I Belgian coast (Van Benedeu) .. 

! 

: Weston-super-Mare (Day) 

, Plymouth (Marine Biol. Assoc.) 

I Brighton (Chililren) ....." 

i Firth of Forth 

I Burford Bank, Irish Sea (Byrne) 

■ Coranna (Seoane) 




Yin-Hays. 



I). 
VI. 12 
VI. 12 
VI. 11 
VI. 12 
VI. 12 
VI. 11 
VI. 11 
VI. 12 
VI. 11 
VI. 11 
VI. 11 
VI, 11 



A. 
12 
12 
11 
12 
12 
11 
11 
12 
11 
11 
11 
11 



G. microps. 



Belgian coast (Van Beneden) 

Weston-super-Mare (Day) .... 
Cuckmere R., Susse.x (Byrne). 

S. Norway (Collett) ....." '. 

Denmark (Day) 



43 


46 


VI. 10 


10 


35 


48 


VI. 10 


10 


42 


47 


VI. 11 


10 


43 


46 


VI. 10 


10 


41 


*7 


VI. 10 


10 


41 


45 


VI. 10 


11 


36 


52 


VI. 10 


10 


35 


42 


VI. 10 


10 


38 


46 


VI. 9 


9 


33 


42 


VI. 9 


9 


33 


44 


VI. 11 


10 


32 


44 


VI. 10 


10 



4i ox FISHES DF THE GENUS GOBIUS. 

As will be seeu from these tables, G. micro'ps has 9 or 10, 
exceptionally 11, rays in the 2nd dorsal and anal, G. minuttos 
having 11 or 12; the number of scales along the lateral line, 
liowever, shows no overlap, not exceeding 52 in G. micro2)S and 
not falling below 61 in G. minvtus. 

I therefore hope I have succeeded in settling the question of 
the correct name of the Hsh so carefully described by Prof. Guitel, 
about which he rightly entei-tained some doubts at the time of 
his observations on its reina.rkable bi-eeding-habits. 

The two forms here discussed have been quite correctly sepa- 
rated and identified by Messrs. Holt and Byrne ; the only point on 
which I cannot agree with them is with regnrd to the existence 
of connec';ing-links, which the examination of a large material 
has failed to disclose. 

In concluding this note, 1 wish to express my indebtedness to 
Px'of. Yves Delage for kindly allowing me to work at the Roscoff 
Laboratory, of which he has the dii'ection. 

Appendix. 

Col. Shepherd, who has devoted much time to the study of 
otoliths, has examined for me those of the two fishes from RoscofF, 
and finds their claim to specific distinction confirmed by the 
difi'erences in this character. 

He has kindly drawn up the following notes, as an appendix to 
my communication : — 

"Under the microscope the otolith of Gohius microps shows as 
a quadrilateral liimp Avith fairly equal sides. Two a,re at a right 
angle, but the angle is rounded off'; a third is bulged out into an 
outward curve, the foui-th forms an indent. The three sides first 
mentioned are plain-edged. 

"The otolith oi Gohinsmivntus^how?^ an irregular quadrilateral 
shape : one side is straight and plain ; the other three sides are 
not so synnnetrically shaped as in G. microps, and are markedly 
scalloped, there being six lobes on the edges of the three sides, 
these lobes not regularly spaced, but of varying size. 

" This would show that the two fishes are different species. 

" The otoliths referred to are in each instance the sagitta." 



ox COl/JLK- PHASES UK TUK HI/ nOKKKJAR. 4") 

EXHIBITIONS AND NOTICES. 

November 29th, 1910. 

Dr. Henry Woodward, F.R.S., Vice-President, 
in the Cliair. 

Dr. William Niooll, of the Lister Institute of Preventive 
Medicine, gave a demonstration of his method for the collection 
of Treaiatodes. 

Dr. R. T. Leiper, F.Z.S., exhibited two pliotograph.s and 
some specimens showing the iSTematode infection known as 
Onchocerciasis in beef imported from Queensland. 



Dr. J. F. Gemmill, M.A., D.Sc, Lecturer on Embi-jology in the 
University of Glasgow, gave an account, illustrated by lantern- 
slides and specimens, of his memoir on " The Development of 
Solaster eiideca Forbes," communicated to the Society by Prof. 
J. Arthur Thomson, F.Z.8. 

This memoir will be published entire in the Society's ' Trans- 
actions ' in due course. 

Mr. D. Seth-Smith, F.Z.S., the Society's Curator of Birds, 
exhibited living examples of the Australian Budgerigar or Un- 
dulated Gi'ixss- Parrakeet (Melopsittamcs tondulatus),iiho\vmg three 
colour-phases. The normal bird was mostly green, with a yellow 
face, dark bai'ring across the occiput and back, and blue on the 
tail-feathers. 

The yellow variety was now common as a cage-bii'd, and had 
been known to occur in a wild state. In it the da,rk pigment had 
disappeared and practically all trace of blue had been eliminated, 
though some spots on the cheek, which in the normal bird weie 
deep indigo-blue, retained a faint bluish tinge. 

The third variety was an extremely I'ai'e one, in which all the 
yellow pigment had gone, leaving the bird almost entirely blue. 
Those parts which in the noniial bird were green, were in this 
variety pale blue, while the face, which was yellow in the normal 
bird, was pure white. 

Blue Budgerigai'S a]:)peared to have been known in Belgium and 
France some twenty-five or thirty years ago, as they were men- 
tioned by Greene in his ' Parrots in Captivity ' (i. 11 7) and othei-s of 
his books, and by Wiener in Cassell's ' Canaries and Ca^e Birds.' 
The variety seemed to have been entirely lest sight of, however, 
in this country at a,uy rate, until M. Pauw^els, a well-known 
Belgian aviculturist, exhibited a pair at a bird-show held at the 
Royal Horticultural Society's Hall at Westminster on November 
25th-2Hth, 1910. This gentleman had several of these liirds, 
which were said to breed ti'ue to type, but to pioduce a, pre- 
ponderance of females. 



46 ox PNEUMO-ENTERITIS IN THE RED GUOUSE. 

Mr. Seth-Smith pointed out that, so far, no systematic breeding 
experiments had been carried out with Budgerigars, but with three 
distinct colour-phases of a free-breeding species to work with, the 
material for some very interesting experimental breeding was at 
hand. 

He acknowledged his indebtedness to M. Pauwels for the loan 
of the blue specimen exhibited at the meeting. 



PAPERS. 

5. On a Possible Cause of Pneumo-enteritis in the Red Grouse 
(Lagopus scotims). By H. B. Fantham, D.Sc, B.A., 
r.z!s., and H. Hammond Smith, M.R.O.S., L.R.C.P., 
F.Z.S. 

[Received Octolter 24, 1910: Read November 29, 1910.] 

The importance of Ooccidiosis as a serious disefi,se of the 
digestive tract of birds has lately been clearly established by 
Fantham in England in the case of young Grouse and Pheasants, 
and by Morse and Hadley in America in Fowls and Turkeys. 

While pursuing our researches at the Frimley Experimental 
Farm belonging to the Grouse Disease Inquiry Committee during 
the summer of 1910, we found that out of 40 Grouse chicks 
hatched, 17 died between the ages of 4 and 6 weeks. These 
birds were examined by both of us and were found to be suffering 
from Ooccidiosis, the parasites (Eimeria (Coccidium) avium) occui*- 
ring especially in the duodenum and cseca. Many of these young 
birds, however, alsDpresented symptoms of pneumonia, consequently 
the lungs, trachea, and bronchi of the birds were most carefully 
examined. The results of our examination were most interesting, 
for we both found coccidian oocysts in the trachea, bronchi, and 
bronchioles. Inside these oocysts the pr-ocesses of formation of 
the four sporobla,sts were sometimes found to be going on. The 
oocysts were probably acquired by the mouth, and a few of them, 
instead of passing directly down the digestive tract, as is usual, 
may have found their way, vid the glottis, into the trachea and 
bronchioles. It is possible that these coccidian cysts in the 
bronchioles would be quite capable of setting up sufficient irri- 
tation to account for the pneumonic symptoms seen in the lung-s 
of these young birds. It would seem, therefore, tha.t the old 
name of pneumo-enteritis, as applied by Mr. Tegetmeier and 
others to one of the diseases that caused mortality in Grouse — a 
view which has met with much criticism — may after all be proved 
to have some foundation in fact. 

References to Literaiiir^. 

Cole, L. J., and Hadley, P. B. (1910).—" Blackhead in Turkeys, 
a Study in Avian Coccidiosis." Bulletin 141, Agric. Expt. 
Station, Rhode Island State College. 

Fantham, H. B. (1910). — "On the Morphology and Life-History 
of Einieria [Goccidium) avium, a Sporozoon causing a Fatal 



ox THE ALIMENTARY TRACT OF CERTAIX BIRDS. 47 

Disease aiiiong young (Irouse." P. Z. 8. 1910, pp. 672-691, 

4 plates. 
Fantham, H. B. (1910). — "Experimental Studies on Avian Cocci- 

diosis, especially in relation to Grouse, Fowls, and Pigeons." 

P. Z. S. 1910, pp. 708-722, 1 plate. 
Kleix, E. (1892).— "The Etiology and Pathology of Grouse 

Disease and Fowl Enteritis." 
Morse, G. B. (1908).— "White Diaxrhcea of Chicks." Circular 

128. U.S. Dept. Agriculture, Bureau of Animal Industry. 
Tegetmeier. W. B.— 'Field,' vol. 104, p. 561 (24/9/04); vol. 105, 

p. 1027 (17/6/05); vol. 106, p. 410 (26/8/05); vol. 107, 

p. 4-65 (24/3/06). 



6. On the Alinientnrv Ti-act of certain Birds and on the 
Mesenteric liclations of tlie Intestinal Loops. B}- Frank 
E. Beddakd, M.A., F.H.S., F.Z.S., Prosector to the 
Society. 

[Received March 1, 1910 : Read Xovember 29, 1910.] 

(Text-figures 9-28.) 

For some time past 1 have been accumulating notes relative to 
the viscera of birds which have died in the Society's Gardens, 
and have paid special attention to the alimentary tract. In the 
following pages I call attention to the intestinal tract of a 
number of birds which either have not been studied or as to 
which my own investigations lead me to disagree with earlier 
statements. 

I have dealt more particularly with such species as have not 
been carefully studied from the point of view of the convolutions 
of the intestine, and am able to call attention to a consideitible 
series of birds. The subject is by no means a new one, dating as 
it does from the accurate though few observations of John 
Hunter. I arrive, however, at rathei' different classificatory 
conclusions from others, and venture therefore to direct the 
attention of the Society not only to the new facts but also to 
certain classificatoiy infei'ences to which these facts point. 

The observations which I lay before the Societ}^ may be 
considered undei' the following headings, viz. : — 

§ Historical Survey, p. 48. 

§ Description of the Intestinal Tract in various Groups of 

Birds, p. 50. 
§ Some General Considerations, p. 86. 
§ The Primitive Form of the Intestine in Birds, p. 86. 
§ The Course of the Evolution of the Gut, p. 87. 
§ The Mutual Affinities of Avian Families judged by the 

Intestinal Convolutions, p. 89. 
§ The Relation.ship between the Gut and the Xature of the 

Food, p. 90. 
§ Summary of Facts relatinc: to the Intestinal Coils of 

Birds, p. 92. 



48 MR. F. E. BEUUAIUJ ON THK 

§ Histurical Survey. 

There is no doubt that the fullest general account of the 
intestinal tra,ct of birds written by the older anatomists is that of 
John Hunter*. He examined and annotated upon more than 
fifty species not wholly though ma,inly British. His observations 
are entirely correct, though not always quite full enough. In many 
cases, however, he has seized the main features of the intestinal 
coils so accurately and sufficiently that but little in the way of 
addition is needed. Thus in the Gallinaceous birds he has 
appreciated the loose ari'angement and absence of fixed loops in 
the postduodenal section of the small intestine and the attachment 
of its terminal I'egion to the duodenum. In the Rails he has 
correctly described the three distinct loops of the jejunal region 
and the attachment of the fii'st and third of these together. In 
the Accipitres the short loop just above the cpeca is described in 
many forms. The peculiaiities of the Parrot intestine are fully 
described. In short. Hunter accomplished a gi'eat deal. 

Cuvier and Duvernoy t distinguished perfectly correctly, as Dr. 
Mitchell has pointed out, the three separate regions in the sma,ll 
intestine of a bird, which I pi-opose to call duodenal, jejunal, 
and ileic loops, and they also indicated the fact that the middle of 
the three loops is frequently folded upon itself, contorted into a 
spiral, or subdivided into several regions. Furtheiiuoi'e. it is 
iema,i-ked (and I find myself in accord with this opinion) that, 
'• le canal intestinal des oiseaux est loin de presenter des differences 
anssi nombreuses, d'une espece, d'un genre ou d'une famille 
a I'autre, que celui des mammiferes." Thereafter follows a 
considerable amount of detail concerning these difFei'ent loops 
in the different groups of birds. For example, the three simple 
loops of the Passerines are referred to in a good many species and 
the spiral arrangement of the middle or jejunal loop is described 
in the Crows. The Picarian birds, Touracou and Cuckoo, are de- 
scribed in such w^ords as to show tha,t they agree completely with 
the Passerines. Cuvier did not, however, as Dr. Mitchell has also 
a,nd quite justly pointed out, delimit the middle region correctly. 
He describes the limits of the third (and last) region of the small 
intestine as indicated sometimes on the side of the middle loop by 
a,n unpaired ctecum (?'. e., Meckel's diverticulum). This is never 
the case, I believe +. 

In his ' Lectures oil Comparative Anatomy,' Sir Everard Home § 
has figured the coils of the small intestine in a number of birds— 
for instance, the Raven, wdiere the spiral of the jejnnum and tlie 
close association of the ileic and duodenal loops are indicated ; the 

* ' Essays and Observations,' ed. bj^ R. Owen, vol. ii., London, 1861. 

t ' Le9o"ns d'Anatomie comparee de Geor.oes Ctivier,' rec. et publ. par G. L. 
Duvernoy, t. iv. 2me partie, Paris. 183o, p. 269 et seq. 

X But see for a possible exfei)tion tbc account of the Tinanious below, p. 52. 

§ ' Lectures on Couijiarative Anatomy,' London. 1814, vol. i. p. 402, vol. ii. 
pis. civ. ~c\-ii. I am indebted to Dr. Mitchell for the exact r.'ferrnce to th's more 
than once misqnotetl work. 



ALIMEXTARV TRACT OF OEIITAIX BrRDS. 49 

Sea-mew, where the spiral is also shown, and it is remarked by the 
author that the turns of the intestine bear a close resemblance to 
those of the Crow, Swan, Goose, " Arclea argala,^' &c. — not a verj 
long series of birds. 

Owen, in describing the Flamingo *, pointed out that the small 
intestines '' were disposed in twentj'-one elliptical spiral con- 
volutions, eleven descending towards tiie rectum and ten returning 
towards the gizzard in the interspaces of the preceding." The 
same anatomist correctly described the three loops in the small 
intestine of the Hornbillt. In the 'Comparative Anatomy and 
Physiology of Vertebrates ' J many more facts are given, most of 
which appear to be quite correct, but all are not quite intelligible 
to mj^self. Not many comparisons are made. The Cuckoo 
is correctly described, but it is not pointed out that it agrees 
with the Hornbill, which bird, indeed, is not referred to in the 
volume. The general prevalence of concentric folds among birds 
with long intestines is noted. The peculiarities of the Galli- 
naceous birds which have no fixed loops except the duodenal are 
appreciated in the description of the Common Fowl. The attach- 
ment of Avhat I term the ileic loop to the gizzard and to the 
duodenal loop is mentioned. 

Dr. Gadow's contributions § to the subject of the present com- 
munication have an importance of their own which is very great. 
But they do not come exactly within the limits of the discussion 
to which I desire here to contribute, since the aim of that 
anatomist was to pourtray the arrangement of the gut within the 
body-cavity and not to delimit only the permanent loops of the 
intestine as formed upon the supporting mesentery. 

The most recent contributions to the subject known to me are by 
Dr. Chalmers Mitchell ||. In these memoirs, the author, in addition 
to discussing some parts of the subject with which I am not 
concerned here, deals with a much larger series of species than 
any previous author and has arranged his observations systema- 
tically, so as to cover most of the existing groups of birds. His 
special object, however, was to trace the various modifications of 
the intestinal tract to what he believed to be a primitive type, to 
arrange them in the form of a phylogenetic tree, and to see how far 
such a tree would agree with or correct conceptions of the phylo- 
genetic ideas regarding birds as a whole. In the course of this 
paper I shall refer to vai'ious points in which my own observa- 
tions do not agree with those of Dr. Mitchell. In my opinion, 
however, Dr. Mitchell's mode of figuring the intestinal tract of 
birds gives an aj^pearance of simplicity which is misleading, with 
the result that birds which are separated by marked characters 

* P. Z. S. 1832, p. 142. t Ihid. 1833, p. 102. 

+ Vol. ii. 1866, p. 167 et seq. 

§ " Vergl. Anatomie des Vevdauungssystemes der Vogel," Jen. Zeit.'sclir. 1881. 
"On the Taxonouiic Value of the Intestinal Convolutions in Birds," P. Z. S. 
1889, p. 305 ; iu Newton's ' Dictionai-y of Birds,' suh voce " Digestive Sy.stem." 

II "On the Intestinal Tract of Birds, Ac," Trans. Linn. Soc. viii. 1903, p. 175; 
and an earlier paper in P. Z. S. 1396, p. 136. 

Proo. Zool. Soc— 1911, No. lY. 4 



50 ME. F. E. BEDDARD ON THE 

are represented as being almost identical. In particular, Dr. 
Mitchell does not always distingnisli between fixed loops, definitely 
formed by a narrow mesentery, and the iri-egular folds into 
which any mobile coil of the intestine may fall when disposed on 
the dissecting-board in Dr. Mitchell's fashion. I shall recur to 
definite instances in the course of this communication. 

1 shall now -proceed to deal with the intestinal tract in a 
number of species of birds. 

§ Description of the Intestinal Tract in various 
G^-oujis of Birds. 

. On opening the abdominal wall of most birds the intestine is 
usually seen to form a rather compacted mass, such as is figured, 
for example, by Dr. Gadow in most of the plates which illustrate 
his original memoir upon the intestinal tract in birds. This 
mass consists of parallel or concentrically arranged loops of intes- 
tine, and in the higher birds, such as a Heron, a Duck, or Stork, is 
very characteristic. This appearance of the gut distinguishes it 
at once from the Mammalian or Reptilian gut, where the intes- 
tine lies laxly within the abdominal cavity. 

This also is the case with all the Struthious birds, whose intes- 
tinal tract at the first glance recalls that of a Mammal. A little 
disturbance of the apparently compact intestinal mass of some 
other birds, as, for instance, the Eagles and Hawks, shows that 
here, too, the intestinal tract is not really much welded together, 
but simply lies pushed close coil to coil, owing to the limited space 
in which it has to be stowed away. In other cases, however, it 
can be easily ascertained by the gentle pulling apart of the 
intestinal coils that the gut is disposed in tightly fixed loops. 

This is the case, for instance, with Ducks, Storks, Penguins, 
and a variety of other genera and families. Inasmuch as the 
lax condition of the small intestine in such a bird as an Ostrich 
recalls that of the Mammalia and Reptiles, and is really like the 
intestinal tract in those Vertebrates, it is to be assumed that this 
condition of the bird's gut is the more piimitive condition and 
that the specialisation into definitely fixed concentrically or 
parallel arranged loops, whether narrower or wider, is an index of 
the higher position of the bird in the series. I shall commence the 
following survey of such new facts as I have to add to the matter 
in hand by dealing with the more primitive groups of birds first. 
Indeed, I have not attempted in this paj)er to map accurately the 
coils in several families of birds where they axe very complicated, 
such as the Stork tribe ; for I am not satisfied as to the relationship 
of the coils in these birds to the more simple intestine of lower 
forms. It is almost entirely with the latter that I deal in the 
present communication to the Society. 

Of the RATiTiE I have examined all the living genera. I fully 
agree with Dr. Mitchell as to the basal position in this group of 



ALIMEXTARY TRACT OF CERTAIN BIRDS. 51 

CasitcD'iitsimd Dromceus, but I find his description defective in one 
particular and the illustration which he gives of Casucunits 
correspondingly inaccurate. It would be inferred from that 
figure * that the gut lay in a single line without any attachment 
between the ileum and duodenum ; that — to use Dr. Mitchell's 
own term — there was no vestige of a supraduodenal loop. The 
existence of this attachment is indicated by him in other cases by 
a cut blood-vessel ; there is no such " short circuit " represented in 
his figure of the Cassowary. Nevertheless, two species of Casso- 
wary which I have dissected, viz. C. australis and G. tvestermanni^ 
show such a connection, which is not, however, associated with 
the formation of an ileic loop distinguishable from the jejunum. 

Nor can I agree with Dr. Mitchell's figure of Apteryx, unless, 
indeed, the species examined by him (^A. manteMl) differs from 
that exaiiiined by myself {A. australis). For I find in the latter 
bird no definite ileic loop, but only an attachment by mesentery 
of the latter part of the ileum to the duodenum. The bird, in 
fact, exactly resembles Gasuarius, Struthio, and the Gallinaceous 
birds in this particular. 

In Rhea aniericana the -intestine is formed upon a plan which 
may be interpreted in one of two ways — one of which is certainly 
not " archicentric " in the sense in which Dr. Mitchell uses the 
word, and the other interpretation hardly justifies the use of the 
word " archicentric." Since, in various other points of structure 
(e. g. less degeneration of wing, syrinx), Rhea is much less 
"Struthious" than Gastoarms, it might be expected that the 
intestinal tract also would be more like that of Carinate birds. 
The accompanying figure (text-fig. 9, p. 52) shows the course of the 
intestine in a female example of Rhea americana, and may be com- 
pared with the figure drawn by Dr. Mitchell t from the intestinal 
ti-act of the same species, with which I do not find myself able to 
agree entirely. Dr. Mitchell, however, is perfectly right in dis- 
tinguishing two loops only in the small intestine, viz., the duodenal 
and another which may or may not be the ileic loop of other birds, 
or " supraduodenal," as it is termed by him. 

This latter loop is wider as well as longer than the duodenal 
loop, and it lies parallel with it as does the ileic loop (nearly 
always) in other birds, and is connected with the duodenal loop by 
the usual ileo-duodenal ligament, which is long and extends nearly 
to the end of the duodenal loop, while it is attaclied along more 
than half of the length of the loop now under consideration. So 
far the facts point towards the interpretation of this loop of the 
small intestine in Rhea as being the homologue of the ileic loop 
of other birds. If this interpretation be correct, then the jejunal 
region or loop will be practically absent and reduced merely to the 
small, tract just where the lower limb of the duodenal loop bends 
round to join the lower limb of the (for the moment) alleged ileic 
loop. There is, I think, nothing intrinsically absurd in this 

* P. Z. S. 1896, p. 140, fig. 3. 

t Trails. Linn. Soc. t. c. p. 183, fig. 3. 

4* 



52 



MR. F. E. BEBDARU OX THE 



suggestion ; it is merely the assumption of the still further reduction, 
of the jejunal region of the gut which is already greatly reduced 
in such birds as Ghunga burmeisteri and Ilouhara macqioee^ii, where 
it is already as short as oi- even shortei' than the ileic loop. There 
is another argument in favour of this interpretation of the two 
well-marked intestinal loops of Jihea which is derived from a 



Text-fig. 9. 




r «s^^ cae. 



Intestinal tract of Shea amevicann. 

Cce. Blind ends of caeca, d. Duodeiial loop. i. Ileic region, id. Ileo-duodenal 
li<;araent. J. Jejunal region In this and the succeeding figures the definite 
loops are marked bj' transverse lines. 

consideration of the Tinamou, Cryptiirns tataupa. In the last- 
mentioned bird the intestin.--)! loops are very remarkable ; they 
are represented in text-fig. 10. There is nothing in particular 
to be said about the dttodenal loop. This is followed by two loops, 
which lie one above the other, the proximal loop lying ventrally 



ALIMENTARY TRACT OF CERTAIN BIRDS. 



53 



to the nioie di.st;)! of the two. They ai'e closely attached to each 
other and to the duodenal by ligament and cannot be freed with- 
out cutting' or tearing. There is, in fact, every reason to regard 
these two loops as a subdivision of the usually single ileic loop. 
Moreover, the ileic loop is occasionally double in other birds ; it is 
distinctly formed of two parallel loops \i\ Anthroj^oides pa7-adisea*. 

Text-fi<r. 10. 




Intestinal tnu:t oi C'rj/ptuyiis tataupa. 
Lettering as in text-fig. 9. 

It is to be noted also that the ventrally situated of the two sub- 
divisions of the presumed ileic loop is attached up to nearly its end 
by ligament to the duodenal loop. There is no case known to me 
among birds wheie the jejunal loop is thus attached. 

Another argument of the saiue kind is to be derived from a 
consideration of the intestinal tract of the Passerine Ixocincla 
craasirostriti. In this BuIIduI, of which I have dissected only one 



* Vide p. 82. 



54 MR. F. E. BEDDARB OX THE 

examole, the tract of tlie small intestine consists of two loops 
only, which are approximately equisized and aie both rather 
wide. Furthermore, the two oval loops of gut are attached to 
each other along their whole length. It will be noted, therefore, 
that these two loops agree in all their characteristics with those 
of other Passerine and many Picarian Birds. But if this be so, it 
follows that the jejunal loop in this Passerine is reduced to the 
verge of disappearance. In any case, whatever be the interpre- 
tation of the several regions of the small intestine in Ixocincla 
crassirostris, it seems to me to be beyond all question that there is 
a very close likeness between its gut and that of Rhea, whether 
the likeness be superficial and due to parallelism of develojDment 
or not. 

Moreover, there is no bird known to me in which the jejunal 
loop has any intimate relations through ligaments with the ileic 
loop — at any rate, to anything like the degree which is exhibited 
in the case of Cryptiirus tataupa, on the view, of course, that the 
jejunal loop is represented. Finally — though naturally it is not 
attempted to lay any very great stress upon this piece of evidence 
— a particular relationship between the Tinamou and Rhea is by 
no means an unreasonable suggestion. 

There is, however, an alternative view to be ta,ken of the intes- 
tinal tract of Rhea americana. It will be observed that Meckel's 
diverticulum lies at about the middle of the lower limb of the 
loop which has been provisionally regarded as the ileic loop ; the 
diverticulum lies nearer to the duodenum, i. e. above the ends of 
the two cseca. This fact would appear perhaps to militate against 
the view that has just been set forth with regard to the intestinal 
tract. For generally, at any rate, Meckel's diverticulum lies on 
the jejunal portion of the intestine and, in fact, at about the middle 
of the length of the entire small intestine. 

But although this may be generally the case in birds, it is by no 
means universally so. In Dendrocygna discolor, for example, I 
find Meckel's diverticulum to be very much nearer to the ileic loop 
than to the duodenal, i. e. to be not by any means in the centre 
of the jejunal region. This is a,lso clearly the case with Gar2Jo- 
coccyx radiatus as shown in Dr. Mitchell's figure *. There is thus 
no absolutely fixed position for Meckel's diverticulum within the 
jejunal region of the gut, though there are no positive facts which 
lead to the inference that this diverticulum may lie within the 
ileic area. If it be held that the existence of the diverticulum 
fixes the jejvinal region of the gut, then the intestinal tract of Rhea 
is simply a slightly further development of that of Casuarius in the 
direction of the Gallinaceous birds and many Picopasseres w^hen 
there is no actual loop formed in the ileic region, but merely an 
attachment by ligament to the duodenal loop. 

The gut of the Ostrich has been described by Dr. Mitchell, as 
well as by othei'S, I have only some small matters to add to the 

* Trans. Linn. Soc. t. c. p. 243, fig. 60. 



ALIMENTARY TRACT OF CERTAIX BIRDS. 05 

account by Dr. Mitcliell in relation to the object of the present 
paper. The duodenal lobe of the example of Struthio molybclo- 
phanes has a lateral branch, as has the species examined by 
Dr. Mitchell. The pancreas extends down the duodenal loop to a 
point rather beyond this lateral diverticulum of the duodenal lobe. 
It does not, however, by a long way reach the end of the loop. It 
does, however, in Apteryx. The attachment of the ileum to the 
duodenal lobe is rather more marked than in Gallinaceous birds 
and much more marked than in Apteryx. The ligamentum ileo- 
duodenale reaches along the duodenal loop to a point beyond the 
posterior termination of the pancreas in that loop. I found no 
fixed loops eitliei' in the moderately long small intestine or in the 
longer colon. 

Text-%. 11. 




Intestinal tract of Talegalla lathami. 
P. Pancreas. Other lettering as in text-fio'. 9. 



The Gallinaceous birds appear to be very uniform in the 
structure of the gut. 1 may take Crax carunculata as a type 
with which the very slight divergences shown by other Galli 
may be compared : the duodenal loop is long and very thick and 
the pancreas extends about halfway down it. The duodenum 
soon narrows to form the jejunal region, which is of considerable 
length and arranged in loose folds which can be straightened 
out and among which are no fixed loops. There is no sharp line 
of demarcation between the jejunal and the ileic region, which 



56 



MR. F. E. BEDDARD ON THE 



later becomes a straight tract of intestine running partly parallel 
with the duodenum and attached to it just for a short spa,ce at the 
root of the duodenum where it (the ileum) bends upon itself 
to join the colon. A second species, a hybrid C. globicera 
and C. hecki^ Avas absolutely identical in all the characteristics 
just given. The desciiption of one species fits the characters of 
the other. 

Text-fio-. 12. 




\ 




^ «*,« 




Intestinal tract of Ortalis mjicauda. 
G. Gall-bladder. Other lettering as in text-figs. 9 & 11. 



In Talegcdla lathami (text-fig. 11, p. 55) the only difierence that 
I could detect was the further extension of the pancreas along the 
duodenal loop, the end of which, howeA^er, it does not reach. 

Among the Phasianidse I have examined a few species, and 
again find no difierences of moment from other Gallinaceous 
birds. In Thcvwinalea {picta and amherstice) the pancreas reaches 
to quite the end of the duodenal loop, and, as in other 
genera, the ileic end of the small intestine (there is, as in other 
forms, no definite ileic loop) is attached to the duodenal loop by 



ALIMENTARY TRACT OF CERTAIN BIRDS. 



57 



t\ not very long ileo-duodeiial ligament. In Eupsychoriyx sonnini * 
the intestine is shorter, but its arrangement is precisely that of 

Text fijr. 13. 




Iiitestinul tract. of Eiiplocaiiius iu/cthemerus, sliowiiig- condition reversed 
from tlie normal. Lettering as in text-figs. 9 & 11. 



* The c8Bca of TSiipKychortt/x sonnini are remarkable in more tlian one svay. When 
the body is opened these tubes are seen to lie in a tightly closed spiral' or rather 
helicoid, producing at first the idea that it is the gut itself which is thus coiled. 
The spiral coiling of the cfeca is not, however, permanent ; they can be uncoiled and 
straightened with the exception of the verj- tip which remains coiled. Each csecum 
moreover, is seen to be covered with a network of bands in which a great deal of fat 
is laid down, and which forms a loosely meshed network with the long axis of the 
interstices corresponding to the long axis of the caecum. Blood-vessels traverse the 
strands and apparently form a corresponding network. I am disposed to compare 
this with the mass of short tubular blind outgrowths from the cKca in the Tinamou, 
Calodromas C Ibis,' 1890, p. 61). A slight tightening of the bands referred to in 
Ezcpsj/cJwrii/a- would cause a bulging of the interstitial tracts and the consequent 
formation of such diverticula. 



58 



BIR. F. E. BEDDARD ON THE 



other Gallinaceous birds, and the pancreas extends quite to the end 
of the duodenal loop. Penelope super ciliaris has also rather a shoi-t 
intestine and the pancreas extends to the end of the duodenal 
loop, thus showing that there is no distinction in this matter 
between the two groups of Gallinaceoiis birds. Ortalis (see text- 
fig. 12, p. 5(5) has also a very simple and short gut. 1 pass by a 
number of other genera that I have examined and which are 

Text-fig. 14. 



7' 




cae. 



Intestinal tract of Podargus cuvieri. 
Letterin"- as before. 



quite like those ali-eady dealt with, to consider a remarkable 
variation shown by Euplocamus nycihemerus. In one specimen 
the typical Gallinaceous arrangement was to be seen ; the calibre 
of the duodenum was much greater than that of the succeeding 
part of the small intestine and the pancreas extended to the 
very end of the duodenal loop. The terminal straight portion 
of the ileum was attached in the usual way by ligament to the 



ALIMENTARY TRACT OF CERTAIN BIRDS. 



59 



duodenal loop. In another specimen (text-fig. 13, p. 57) the dis- 
position of the jejunal and ileic regions was exactly reversed*. 
The duodenal loop passed immediately into a straight descending 
limb bent sharply upon itself at its lower extremity, and 
then passed into a laxly coiled and rather long section of gut 
unattached anywhere to the duodenal loop and ended eventually 
in the colon. The laxly coiled region of the gut lay to the left 
side instead of to the right, and there was, in fact, in this 
individual an exact reverse of normal conditions, 

Text-fii?. 15. 




Intestinal tract of Gymnorhina leuconota. 
Letteriiis as before. 



It is thus evident that the intestinal tract of the Gallinaceous 
birds is very uniform throughout the group and that it is con- 
stituted upon a primitive plan which is very little, if at all, in 
advance of that which characterises Apteryx among the Stru- 
thious birds. The only difference is, indeed, that the ileo-duodenal 
ligament is longer among the Gallinaceous birds — that more of 
the ileum is attached to the duodenum. But this condition is 
more than paralleled by Struthio^ where, as already mentioned, 

* I compare these later (p. 79) with the normal conditions occurring in Fraiercula. 



60 MR. F. E. BEDDARD ON THE 

a considerable tract of ileum is attached by the ileo-duodenal 
ligament. 

The PiCARiAN Birds, Cuckoos, Touracous, and Passerines 
really form one grouj) so far as their intestinal convolutions go. It 
is impossible, as I think, to distinguish between the Picarian 
Podargus cuvieri * and the Passerine Gymnorhhia lenconota, 
which may be compared a,nd cannot be contrasted in the accom- 
panying figures (text-figs, 14, 15, pp. 58 & 59). The salient 
features in these two cases appear to me to be the great width 
of the ileic loop (its extreme narrowness in e. g. the Parrots places 
them at the op]30site end of the series) and the close mesenteric 
connection by the ileo-duodenal ligament of the two loops in 
question. The great width of the ileic loop in Nyctidromus, 
Cypselus^ and Trogon can be inferred from Dr. Mitchell's figures T, 
though he does not, except in the case of Cypsehis^ identify the 
loop. These figures suggest undoubtedly the primitive gnt of a 
Ratite or Gallinaceous bird ; j^erhaps they are compai'able with 
MelaneiyesX. 

Furthermore, the total absence of specialised loops in the 
middle region of the small intestine is to be noted. Mitchell, as 
well as his predecessors in this field, has commented upon the 
spiral arrangement in certain Passeres, and has remai'keil upon 
the tendency to a spiiul even where there is no a,ctual regular 
spiral formation. This affects the middle or jejunal loop, and is 
greater in the Raven than in any other bird which Mitchell has 
described or I have examined. I found in that bird a spiral of 
no 'less than nine double turns, whereas Mitchell has figured much 
fewer in Corvus capellanus. A complete spiral of this kind is, 
however, not common among the Passeres. Besides the Crov/ 
tribe I know it only in the Tanager, Euphonia v'olacea. The 
tendency to a spiral I have observed in many Passeres, among 
which I may mention anumbei'of Birds-of- Paradise which I have 
lately had the opportunity of studying : these ai'e Diphyllodes 
hunsteini, Paradisornis rudolplii^ Paradisea raggiaria. 

It seems to be universal or nearly so for the ileo-duodenal 
ligament to connect those two loops of the intestine along their 
whole lengths, and also for the pancreas to extend up to the very 
end of the duodena,! loop. I have found both these characters 
to exist in Ixocincla crassirost7-is §, Sycalis flaveola, Eaplionia 
violacea, Gracidus religiosus, Buaros cylindrica, Ttbrdus migra- 
torius, Ptilonorhynchus violaceus, Cassidix oryzivora, and the first- 
named character in a number of other genera of which I happen 
to have no note as to the pancreas. Both these anatomical 
features seem likely to be characteristic of the Picopasseres 
generally, even if not univei'saljy found among the members of 
that order of Birds. 

* 1 have examined two specimens of this hlrd. 

t Trans. Linn. ^Soc. torn. cit. figs. 08, 69, 70. 

X V. infra, p. 63. 

§ A peculiarity of the gut of this Passerine has been already referred to, v. p. 53. 



ALIMENTARY- TRACT OF CERTAIN BIRDS. 61 

I have not met with many divergences among the Picopasseres 
from the typical structure. 

One of the most abnormal types — if not the most abnormal — 
among the Picopasseres is the Ground Hornbill, Bucorvus ahys- 
sinicus. The duodenal loop is longish and the pancreas extends 
nearly to its end. It is perfectly free fi-om the ileic loop, which 
is longer than it. The ileic loop, moreov^er, is indented at its free 
extremity and tlius shows signs of being bent over upon itself. 
It is also considerably longer than the duodensil loop. As in 
other Picopasseres, the jejunal loop is more or less divided into 
two, and the distal loop of these two is attached to the outgoing 
limb of the ileic loop, which on its way to the colon is looped once 
in a way precisely like that shown funong tlie Accipitres and in 
some other birds. These facts are particularly intei-esting, because 
they confirm current opinion as to the anatomical likenesses 
between the Hornbills and the Hoopoe. It is plain from 
Dr. Mitchell's figure * of the intestinal tract of that bird that 
Upupa epops agrees with Bucorvus in a number of the characters 
to which I have referred above. He figures the two loops of the 
middle part of the intestine and the small '"supracpecal" loop, 
which latter is so characteristic a feature of Bucorvus as compai'ed 
with other Picarian birds. He does not, however, advert to this 
loop by that name or compare it with the " kink" which he found 
in the Accipitres of both the Old and New World. Nor does he 
indicate a mesenteric attachment between the jejunal and ileic loops 
in Ujnipa such as I find in Bucomus. It is impossible, moreover, 
to be certain from Mitchell's figure how far the ileic and duodenal 
loops are connected. Theii' entire mutual freedom in Bucorvtts is 
an uncommon feature. Although Dr. Mitchell happens, as I think, 
to be wrong in remarking that the cha,racter of the gut does not 
unite the Hoopoes and Hornbills closelj^ he was perfectly right 
in making that statement from the facts before him. This is 
a further example of the difficulty of arriving at sound cla^si- 
ficatory conclusions without an exhaustive knowledge of the 
facts. 

I have lately had the opportunity of examining the gut of 
Upupa, and can add something to the account given by Mitchell. 
It is a rather move abnormal member of the Picopasserine group 
than I had supposed. In my specimen there was no supracpecal 
kink. The duodenal loop was very wide (as Mitchell has 
remarked) and rather irregular in outline at its end, suggesting, 
therefore, a commencing spiral as in irypagus and Cathartes 
— a fact which may be of some significance. The duodenal 
loop is larger tha.n the ileic — precisely the reverse condition 
obtaining in Bucorvus. And while in Bucorvus there is no 
ileo-duodenal ligament, there is a short one in Upupa not 
nearly so extensive as in Picopasseres generally, and thus 
bridging over the gap between Bucorvus and its allies. 

* Trans. Linn. Soc. i.e. p. 247, fig. 65. 



62 



MR. F. E. BEDDARD ON THE 



The jejunum has certainly two definite loops and thus, 
agreeing with Bucorvus, differs from other Picopasseres. The 
pancreas in a most abnormal fashion extends into the first of 
these and is perhaps responsible for its formation. 

It is, of course, possible that the intestinal tract of Melanerpes 
superciliaris differs from that of other Picidje, But if it agree 
with that of the three species reported on by Dr. Mitchell, 
then I find myself in total disagreement with that writer as to 

Text-fig. 16. 




d. 



Intestinal tract of Melanerpes superciliaris. 
Lettering as before. 



the relationships of the Woodpeckers. He observes of the Picidse 
that " the conformation of the gut is in every important respect 
similar to that found in Megcdcema." I have not dissected 
Megalcema for the purposes of the present communication, but 
I have examined three species of Toucatis, of which family 
(Rhamphastidse) Dr. Mitchell remarks that '• Meckel's tract 
and the short rectum do not dififer from the form found in 
Megalcema." Now in Megaloima asiatica, as is plainly shown in 



ALIMEXTARY TRACT OF CERIVI-V BIRDS. 



63 



Dr. Mitchell's figure *, there is a distinct ileic loop, which is wide 
as in Picopasseres genendly. In the Toucans Aulacorhamphus 
sidcaius, Rhanvphastos ariel, Eh. carinatiis, there is also a perfectly 
distinct ileic loop a little less distinct than in some Picopasseres, 
but still distinct. 

In Melanerpes superciliaris the only properly marked intestinal 
loop is the duodena], down to the very end of which extends 
the pancreas. There is absolutely no ileic loop, as is shown in 
the figure (text-fig. 16). The jejunum simply passes forward 
and is connected by ligaments with the gizzard and with the 
duodenum quite far from the free end of the loop ; it then turns 
abruptly backwards, passing straight to the cloaca. 

The arrangement of the gut is, in fact, precisely that of the 
Gallinaceous birds, though, of course, the gut is shorter than that 
of most. 

Text-fig. 17. 





id. 



\ 




..^^ 



I. 



Intestinal tract of Gecinus viridis. 
Lettering as before. 



There is, indeed, no great disparity in length between the gvit 
of Melanerpes and that of an equisized Gallinaceous bird, such 
as Coturnix chinensis. In view of the primitive nature of the 
palate as urged by Huxley and Parker, though not held by some 

* Trnns. Linn, Soo. torn. rit. tig'. 71, p. 253. 



64 MR. F, E. BEDDARD ON THE 

others, the existence of a primitive type of gut is not without 
interest. There is in any case no doubt about their difference 
in this respect from their supposed allies the Rhamphastidee, 
though it remains to be seen whether they are like Picarian 
birds of any other groups. These facts and considerations gain 
additional significance from the quite similar intestinal tract of 
Geciuus viridis (see text-fig. 17, p. 63). 

Opinions undoubtedly differ as to the geiieric subdivisions of the 
family Alcedinida?, but Alcedo ispida and Halcyon sancta have 
been placed in different genera — whether Halcyon or Sattropatis. 
The intestinal tract is, however, rather different in these two 
species, though one may be considered to be au exaggeration of 
the othei'. The simpler of the two is that of Alcedo ispida. 
In this Kingfisher the duodenal loop is free from the ileic for 
at least the greater part, and thus conti^asts with most other 
Picarian birds. The jejunal region lies in a short spiral; but 
this spiral is not a permanent structure. It can be easily 
disarranged and spread out into an irregularly shaped loop. 
There is, in fact, no mesenteric connection between the circles of 
the spiral. The ileic loop is large, wide, and somewhat irregular, 
fully as long as the duodenal loop. The spiral of the jejunal 
I'egion is, it should be added, quite a short one with only two 
complete turns. 

In Halcyon sancta there are differences in nearly all of these 
features. The duodenal loop is, however, the same ; it is a simple 
loop, not particularly wide, and the pancreas extends along it 
quite to its tree end. I omitted to make any notes about the 
pancreas of H. vagans. The jejunal region of Halcyon sancta forms 
a spiral of eight limbs, and is thus, in the first place, much more 
complex than that of Alcedo ispida. In the second place, this 
spiral is fixed, and is a perfectly permanent sti-ucture which 
cannot be unwrapped without tearing the connecting sheets of 
mesentery. These are two important differences from the spiral 
found in Alcedo ispida and are, indeed, much greater differences 
than are known to me to exist between two species of any other 
genus. The condition of the jejunal section of the small 
intestine does not, however, exhaust the diffei'ences which even- 
tually distinguish these two species of Kingfishei's. 

The ileic loop is, as in the last species, quite free from the 
duodenal ; there is no ileo-duodenal ligament, except perhaps at 
the very base of the otherwise mutually free loops. The loop 
is, however, double, as it is, for example, in Grits japo'iiicus *, 
and as is shown in text-figure 18. Of these two loops, the 
proximal is the larger and is wide and somewhat irregular in 
form, and of about the same length as the duodenal. On the 
whole, it may, as I think, be admitted that the difference which 
the alimentary tract of this Picarian bird shows from that of 
other Picarian birds is actually greater than that which exists 

* Vide p. 82. 



ALIMENTARY TRACT OF CERTAIN BIRDS. 



65 



between the Limicoline Recurvirostra and such a Passerine as 
Euphonia — in this case, two entirely different groups of birds. 

Among the OucuLiIhave examined only Euclynamis orientalis, 
which is one of the genera which does not seem to have been 
examined by previous writers from the present point of view. 

Text-fig. 18. 




Intestinal tract of Grnsjapoiiicus. 
Lettering as before. 



So far as I am able to say from the examination of this one type, 
Miss Marshall's figure * of Geococcyx ccdiforniavMS is a better 
representation of the characters of this group than that given by 

* " Studies in Avian Anatomy. — II.," Trans. Texas Ac. Sci. ix. 1906, pi. ii. 
fis?. 22. 

Proc. Zool. Soc— 1911, Xo. V. 5 



66 MR. F. E. BEDDARD OX THE 

Dr. Mitchell *. For the jejunal region possesses no definite loops 
and the gut, as a whole, is entirely like that of the majority of 
the Picopasseres. In Eudynamis orientalis the pancreas extends 
to the very end of the duodenal loop ; the ileo-duodenal ligament 
also extends in its attachment to nearly the end of the duodenal 
loop. The ileic loop is, however, longer than the duodenal. 
In the jejunal region there are no specialised loops, and this part 
of the gut shows indications of a spiral arrangement. 

Of the MusophagidjB I have examined two examples of 
T'uracus macrorhynchus and one of T. bujf'oiii. The two 
individuals of T. macrorhynchus agreed absolutely ; the gut of 
T. huffo7ii differed very slightly from that of its congener. In 
Turacus macrorhynchus the duodenal and ileic loops were closely 
connected throughout their whole length by an ileo-duodenal 
ligament. The loops were moderately wide and of equal length 
or very nearly so. The jejunal loop is a little longer than either 
of the others and has a slight tendency to a spiral ; Avhen 
straightened out forcibly it lies in a Y shape with some slight 
rotation, a,s Hunter has figured in the case of Scythrops 7iOvce- 
hollandice t. The pancreas, it should be observed, extends down 
to the very end of the duodenal loop. The only diff"erence that 
I could detect in Turacus huff'oni is that the ileic loop is rather 
longer than the duodenal. It is clear that the intestinal tract 
of these birds is precisely like that of the Cuckoos and of the 
majority of the Picopasseres. 

Of the group Hemipodii or Turnicbs I have dissected two 
examples of the species Turnix varia. They were quite identical 
in the convolutions of the intestinal tract. The duodenal and 
the ileic loops were attached up to the end or very neai'ly so by an 
ileo-duodenal ligament. The loops were also fairly broad. The 
jejunal region of the gut lying between these two loops at either 
extremity of the canal was formed of a single loop, which had a 
tendency to twist itself into apparently two loops ; but, without 
tearing or in any way interfering with the mesentery, this part 
of the gut could be moulded into the characteristic Passer-ine 
plan, as is shown in the accompanying figure (text-fig. 19). The 
pancreas extends right to the end of the duodenal loop. It is 
obvious from what has been said that this bird has a typically 
Passerine gut. It has not the faintest likeness to any Gallinaceous 
bird. Its likeness to many Passerines is shown by the fact that 
the pancreas extends down to the very end of the duodenal loop, 
and also by the breadth and connection up to the very end or 
nearly so of the ileic and duodenal loops as well as by the slightly 
spiral, and limited, jejunal loop. 

AcciPiTRES. — Among the Accipitrine birds which have not been 
examined by Dr. Mitchell I have dissected the Harpy Eagle, jyarjt??/- 
haliaetus coronatus. The duodenal loop is moderately wide and the 
pancreas does not extend far down it. The ileic loop is also fairly 

* Loc. eit. p. 242, np. 60. 

t ' Essays and Observations,' vol. ii. p. 286. 



ALTMICNTARV TIlACT OF CICIfTAIX RiriDR. Ul 

broad, <ar>d contrnsts vci-y markedly with tlie duodeiinl loop, on 
account of the small calibre of the intestinal canal in this region ; 
the duodenum is particularly wide and tapers off gradually 
towards the jejunum, the greater part of Avhich is also very 
narrow ; the tube is, in fact, quite as narrow here as is the ileum. 
The contrast between the duodenum and the comnaencement of 
the jejunum on the one hand, and the rest of the jejunum and the 
ileum on the other hand, is remarkably like that seen in the 
Gallinaceous birds, e. g., Cr«,r (see p. 55). The ileic loop is as long 
as the duodenal loop or very nearly so. It is connected to it by a 
ver}- short ileo-duodenal ligament, which leaves almost the whole of 

Text-fisr. 19. 




Intestinal tract of Tnrnix varia. 
Lettering as before. 



both loops free of each other. The jejunal region is of considerable 
length, and lies loosely and irregularly folded in the body-cavity for 
the most part. The commencement of this part of the intestine, 
however, where it joins the duodenum is not only, as already 
mentioned, of greater calibre than the rest, but is fixed in a short 
and wide and therefore not very well-marked loop. The rest of 
this section of the intestine lies loosely like the Mammalian small 
intestine or the jejunum in Gallinaceous birds — that is to say, it 
has no fixed loops, but can be passed in a straight line between 
the fingers without tearing or distorting the mesentery which 

5* 



68 MR. F. E. BEDDARD ON THE 

supports it. Finally, between the ileic loop and the colon is a well- 
marked supracsecal fold, found in all Accipitrine birds and in some 
others. Of other Falconidse I have examined the following species, 
which are not referred to in the memoir by Dr. Mitchell, 
viz. Gerayioaetus melanoleucus, Astur palunibarius, Tinnunculiis 
alaudarius, Astur approximans, as well as one or two species that 
are referred to by him. 

In Astur paluinharius and A. approximans the gut shows no 
marked differences from that of Ilarpyhaliaetus, which I have 
taken as the type for this group. The pancreas extends but a short 
way along the duodenal loop, and the ileic loop is attached to the 
duodenal by a short ligament only at the base. The supracsecal 
loop is present, and the jejunum cannot be said to possess any 
pronounced folds indejaendent of each other. As it lay in the 
body I noticed the formation of spirals in this region of the gut 
in A. pahtmbarius ; but these were not permanent formations 
as in Recurvirostra avocetta. The Avliole of the jejunum could be 
straightened out bit by bit, the most pronounced fold, close to the 
duodenum, being large and wide and hardly comparable to the 
definite folds in the jejunum of more specialised birds, such as 
the Psittaci. Geratioaetus melanoleucus shows again no salient 
differences ; the supracfecal fold, however, is nearly as long as the 
ileic loop. 

I agree with Dr. Mitchell in regarding the gut of Falco 
as being aberrant when compared with that of other Hawks and 
Eagles. The duodenal loop is, as he has said, irregular in form. 
I may add that the pancreas extends a good deal further down the 
duodenal than in the other Accipitres hitherto dealt with in the 
present communication. Even the ileo-duodenal ligament is a 
little more extensive than it is in Astur &c. 

In Tinnunculus alaudarius there is a,n exaggeration of the 
" abnormality " of the duodenal lobe, which is almost bent upon 
itself in a spiral fashion. In this Hawk the greater part of the 
jejunum is disposed in a temporary spiral coil ; bvit the first part 
of the jejunum is in the form of a single loop, which is comparable 
to that figured by Mitchell in Falco *, and which I have described 
above in Harpyhaliaeius. 

Spizaetus hellicosus (see text-fig. 20)is another species upon which 
Dr. Mitchell had not the opportunity of reporting. It agrees with 
other Accipitres in its general characters, but there are some minor 
points of difference. Thus, the disposal of the jejunal is exactly 
what we find in Harpyhallaetus coronatxis. This region of the 
gut commences with a very wide stiff loop and then passes into a 
loosely folded length of tube. The ileic loop is rather longer than 
the duodenal, and the supracsecal kink is developed into a loop 
nearly as long, the two together reminding us of the double ileic 
loop of the Cranes and even the Tinamous. 

* 111 both of two examples of Falco peregrinus I have not seen a marked loop 
correKponding to this. The jejnmim lay entirely or mostly in a rough spiral, which 
could be arranged in an irregular circular fold. 



ALIMENTARY TRACT OF CERTAIN BIRDS. 



69 



I have dissected one example of the New World Vultin-e, 
Gypagvbs papa, whose intestinal tract I am able to compare with 
that of Cathartes aura described by Mitchell. In view of the 
fact that the American Vultures have been regarded by many 
systematists as being quite remote in their affinities from the Old 
World Vultures, indeed from the Accipitres generally, it is 
very important to bring together all contributions that are possible 
towards settling this vexed question. The duodenal loop differs 
from that of Cathartes anrl is, indeed, quite like that of Falco 
feldeggi figured by Mitchell*. The distal extremity is folded over 

Text-fig. 20. 




Inteslinal tract of Spizaetus bellicosus. 
Lettering as before. 



upon itself in a fashion that does not appear to occur among 
Picarian birds. The jejunal loop is arranged in a spiral fashion like 
many birds, including, however, Tinnunculus, in which Gypagus 
appears to difier from Cathartes. The ileic loop is simple. The 
kink so characteristic of Accipitres (but also found in other birds, 
including Bucorvus) above the position of the caeca in other birds 
is present as in Cathartes. It is not, in fact, possible to locate 
Gypagus definitely in the sj^stem. 

I have also examined Cathartes aura (see text-fig. 21), and in 
most matters I am able to confirm Mitchell, as will have been 

* Trans. Linn. Sec. t. c. fig. 33, p. 211. 



70 



MR. t\ E. BEDUARD OX THE 



inferred from what has been ah'eady said about Gypagus. The 
spiral of the duodenum makes rather more than a complete circle 
and its limbs are fixed together by mesenteries ; the gut does not 
simply lie in a spiral. The loop which is lettered " I " in Mitchell's 
figure is broader and not so narrow accoi-ding to my observations, 
but better marked than in Gypagus. As to the following portion 
of the small intestine, I do not agree in detail with Mitchell. 

Text-fiff. 21. 




Intestinal tract of Cathartes aura. 
Lettering as before. 



The two loops forming it run close together, so that it is long 
and narrow, and not broad as figui'ed by Mitchell. There is a 
tendency to form a rough kind of spiral not nearly so marked as 
in Gypagus. Tlie ileic loop is only attached to the duodenal by 
mesenteiy at its very base. The supraCsecal loop is more marked 
than in Gypagus. 



ALIMENTARY TRACT OF CERTAIN BIRDS. 



1 



Striges. — Among the Owls, I have examined Scotopella houvieri, 
which has not yet been investigated from the present point of 
view. The duodenal loop is widish and there is no trace of an 
ileo-duodenal ligament — in fact, the ileic loop lay on the right side 
of the body. The ileic loop is quite simple and is about as long as 
the duodenal. The jejunum shows three quite distinct loops ; 
the fii'st of these, i. e. that immediately following upon the 
duodenum, is wider than, but of about the same length as, 
another short loop which immediately follows it. A third loop of 
greater length has a distinct hint of spiral twisting. In Syrnium 
ahcco, Asio otus (see text-fig. 22), Strix perlata, JVmox hoohook, 
Bubo maxinncs, B. virginianus, B. maculosas, B. chierascens, and 
Strix flammea, the ileic and duodenal loops are connected by a 
ligament which extends about halfway along the former loop. 
The diflerence is rather extraordinary in the matter of this 
ligament between Scotopelia and other genera, and it is, of 
course, possible that we have to do with an individual variation 
of Scotopelia houvieri. 

Text-fi"-. 22. 




Intestinal tract of Asio oli 
Lettering as before. 



The division of the jejunal tract of the small intestine into 
separate loops is not always well marked. In Babo maculosus, for 
example, there are no fixed folds whatever between the duodenal 
and ileic loops; the whole of the jejunal region is like that of 
the Gallinaceous birds or the Mammalia, and can be passed through 
the fingers in a straight line without rupturing or even straining 
the mesentery. The same statement applies to Bubo cinerascens, of 
which species I have dissected two examples. I noticed here that 
the undisturbed jejunum lay in slight spiral coils ; but these were 



72 MR. F. E. BEDDARD ON THE 

in no way permanent structures, but could be readily pulled out 
straight. In Btiho virginianus the greater part of the jejunum 
lies in the same way in loose movable coils ; but there is a hint 
of a fixed loop — wide and shallow — at the commencement of the 
jejunum. Bubo capensis (I have seen two examples) and 
B. maximios were quite like B. maculosus and B. cinerascens, a 
slight and quite unfixed spiral being particularly noticeable in the 
case of the first-mentioned species. 

I quite agree with Dr. Mitchell that Strix flammea has a 
jejunum which may be regarded as archicentric. I cannot, 
however, understand why Dr. Mitchell should emphasise the 
archaic character of the gut of this Owl by terming it "remark- 
ably archicentric " and by figuring a state of afiairs * which is not 
at all archicentric. For, in his figure of Strix flammea, there is 
correctly represented a well-marked " supraduodenal fold " — or 
ileic loop, as I prefer to call it — and a definite loop in the jejunal 
region. The latter I did not find in that species of Strix ; but I 
am not at all disposed to dispute the accuracy of Dr. Mitchell's 
figure. For in Strix perlataf the jejunal fold is disposed in 
three more or less equisized and not very close loops ; but still 
they appear to be definite loops, and the jejunum is not merely 
a loose coil as in Bubo. This species is therefore not at all 
archicentric. 

The conditions seen in Strix perlata seem to me to be a slight 
exaggera,tion of those which I noticed in Strix punctatissima. 
In the latter Owl the jejunum is formed by a tube which lies 
in the undisturbed intestine as a spiral. It can be smoothed 
out without tearing any mesenteric connections into the not 
circular but rather W-shaped coil so characteristic of Passerine 
and many Picarian birds ; the rest of the small intestine is of 
less calibre and becomes suddenly so ; it again lies in the body 
in a spiral fashion, but can be smoothed out in the same way 
into a broad but rather irregular ileic loop. The ligamentum ileo- 
duodenale extends nearly to the end of this and is attached to 
about halfway down the duodenal loop. The pancreas extends 
for rather more than halfway down the duodenal loop. The 
gut of this genus is considerably shorter than in, for instance, 
Bubo, and is to be contrasted by its stiffness with the lower coils 
of the latter. 

Athene noctua, being a small species, might be expected to show 
those diflferences from other Owls which are often met with in 
comparing small species with larger allies. As a matter of fact, 
the jejunal region of the gut is comparatively short and shows 
no trace of any fixed loop such as occurs in some other Owls. 
In this species the pancreas extends to the very end of the 
duodenal loop; in a species of Ciccaba, in Strix flammea. Asia 
otus^ Ninox hoobook, Bubo macidostcs, B. cinerascens^ B. virgini- 
anvjS, and some other Owls, the pancreas does not extend so far 

* Trans. Linn. Soc. t. c. fig-. 66, p. 21.8. 

t It is not cpi'tiiiu how far tlu'Sf alleged species of 8trLv have that value. 



ALIMENTARY TRACT OF CERTAIX BIRDS. 



73 



clown the duoilenal loop, but it is longer than in tlie Accipitres. 
This fact, indeed, and the rather greater extension of the 
ligamentum ileo-duodenale, is the chief difference that distin- 
guishes the Owls from the Accipitres, the general plan of the gut 
in these two subdivisions of birds being otherwise similar in many 
ways. 

The groups that have been hitherto considered, viz. the Ratitfe, 
Galli, Hemipodii, Picopasseres, Ouculi, Musophagi, Accipitres, and 
Striges, agree with each other in that the jejunal region of the gut, 
though it may vary gi'eatly in length, is never thrown into much 
marked fixed loops, such as those which charactei-ise the groups of 
birds that remain to be dealt with. There is, indeed, the commence- 
ment of the formation of such loops to be seen in the Accipitres 
and Striges ; but they do not arrive at the perfection and com- 
plexity of interconnection which is exhibited in the i-emaining 
families of Birds. Among the latter, however, with which I shall 
proceed immediately to deal, there are species and even genera 
which show the simpler conditions of the jejunum that characterise 
the families of Birds already dealt with — for instance, in Phcvianus 
among the Limicolae and in the Bustards and Cariamida?. In the 
latter the simple conditions look like reduction ; while in Pluvianus 
we may have to deal with an archaic representative of its family 
which has not yet cast off the comparatively primitive type of 
gut. 

Alectorides. — The Bustards are an example of a well-marked 
family of birds which show a great uniformity in their intestinal 
tract. The species which I have myself examined are Eupodotis 
australis and Houhara macqueeni, and they evidently agree with 
Otis tarda as described by Mitchell *. In Houbara macqueeni the 
duodenal loop is attached to the ileic by a ligament which extends 
to the very end of the former ; the ileic loop is considerably 
longer than the duodenal. The pancreas extends as far as the 
very end of the duodenal loop. The jejunal region is formed of a 
single fixed loop, which is not quite so narrow as is depicted in 
Otis tarda. Eiqjodotis australis (see text-fig. 23) has an intestine 
which is so like that of Houbara that I can find no fresh terms in 
which to describe it. Of birds admitted to be possibly allied to the 
Bustards, that which most closely resembles the two genera just 
referred to is Chunga hurmeisteri. The resemblance, howevei-, 
does not quite reach, though it> very nearly approaches, identity. 
In this bird the duodenal loop is, as in the Bustards, shorter than 
the ileic. But the ileo-duodenal ligament stops about halfway 
along the duodenal loop, though extending further along the ileic. 
A point of likeness to the Bustards is the extension of the 
pancreas to the end of the duodenal loop. The jejunal region of 
the gut is also like that of the Bustards, in that it consists of but 
one loop which occupies the whole region, of which, in fact, this 
section of the intestine solely consists. Here we have an obvious 

* Trans. Linn. 8oc. t. c. p. 226, fig. 45. 



74 



ME. F. E. B1:L)DARD OX THE 



likeness to the Bustards. The loop differs, however, in being 
considerably wider than it is in the Bustards. These birds form 
together a part of Mr. Sclater's Order Alectorides, and, as they 
obviously agree together very closely in the characters of the gut, 
I deal with these provisionally under that name ; for there is, at 
any rate, no very general agreement as to their position among 
I'elated groups and their affinities with each other. 



Text-fiff. 23. 




Intestinal tract of Eupodotis australis. 
Lettering as before. 

Among the Limicol/E, with which, as 1 think, the Gulls and 
Terns are obviously to be placed, there are several variations to 
be seen in the coils of the alimentary tract. The most pi-imitive 
form of the alimentary tract known to me among those birds is 
shown in the case of Pluvianus cp.gyptius, for leasons which I 
shall indicate after describing the facts. The duodenal loop is 
fairly wide and the pancreas extends back to the very, end of that 
loop. The jejunum is not definitely distinguishable from the 
ileum, but the whole length of the small intestine, before it bends 
upon itself to form the straight region which bears the small 
and Passerine caeca, is loosely disposed as in Gallinaceous birds. 
The last part of this jejuno-ileic region runs, as in Gallinaceous 



ALIMENTARY TKACT OF CERTAIN' BIRDS. 



75 



birds, parallel with the daodeual loop and is attached to it l»y an 
ileo-duodenal ligament, which extends nearly to the end of the 
duodenal loop. It is, as I think, possible to interpret this intes- 
tinal tiact in only one way, and that is as follows. 

It has been compared with that of a Gallinaceous 1)ird, and this 
is really tantamount to saying that in the coils of the intestinal 
tract Fluvianits presents us with archaic characters. After the 
duodenal loop there is no mai-ked differentiation of the gut 

Text-fig. 24. 




Intestinal tract of Plicviaitus agi/piius. 
Lettering as before. 



into special loops at all. There is, however, as it appears to me, 
an indication of an advance upon the condition of the gut which 
characterises the Gallinaceous birds and in the direction of some 
other Limicolous birds. In the Gallinaceous birds the distal 
extremity of the small intestine is straightened out, but it is 
relatively only a small part of the jejuno- ileum which is thus 



76 



MR. F. E. BEBDAKD ON THE 



diiferentiated from the much longer coiled region. Now, in 
Pluvifinus the distal portion of the jejuno-ileum can be cle rigeur 
divided off fi-om the point lettered a in the figure (text-fig. 24), 
after which point the intestine exhibits no resting in loose folds, 
but passes in a broad curve to the point where it is attached 
by a ligament to the duodenum. We have, in fact, here the 
commencing separation from the jejunal region of a very wide 
ileic loop. There is, in fact, a close similarity with the gut 
of Melanerpes *. A slight alteration in the gut of both of these 

Text fiff. 25. 




Intestinal tract of CEdicnemus scolopax. 
Letterincj as before. 



birds leads to that of many Picopasseres, where the ileic* loop is 
more definitely marked oif from the jejunal but remains very 
broad. From this type, moreover, can be readily deduced the 
plan of intestinal coiling which is found in some other Limi- 
colous birds which I have examined. One of the simplest of these 
is Sarciopho7-us pectoralis, in which the ileic loop is attached to 
the duodenal for nearly its whole length by the usual ligament 
and is also a wide loop. The jejunal has no fixed loops, but lies 

* Vide p. 62. 



ALIMENTARY TRACT OF CERTAIX BIRDS. 77 

in rather stiff coils, which approach a spiral ; there are about three 
of these, so that the gut is not long. It is quite difficult to 
differentiate this gut from that of most Picopasseres. OMicnemus 
(CE. SGolopax and CE. grallarius) hardly differs from Sarciophorus. 
Of this genus I may, in the first place, i^emark that the pancreas 
extends back to the very end of the duodenal loop. The attachment 
of this loop to the ileic is as in Pluvianus. The ileic loop, more- 
over, at any rate in CEdicneinics grallarius^ is very wide, another 
point of likeness to Pkoviaiius and, indef^d, to other Limicolous 
birds that will be mentioned presently ; indeed, in QiJ. scolopax 
(see text-fig. 2.5) a separate ileic loop can hardly be defined — this 
character, in fact, is of the group. In both species of the genus 
(Edicnemus that I have examined the jejunum lies more or less 
in a spiral, which is most marked in CE. gr^allarius, though it is 
only a short spiral of one complete turn even in that species. 
It is particularly to be noted that this spiral, like that of Passerine 
birds, is not a permanent spiral, but that it can be pulled out to 
form an irregular circle without tearing any mesenteries. The 
genera which have just been dealt with are, in fact, not far 
removed from the common ground-plan, and the steps of differenti- 
ation are quite as is found in the great division of the Picopasseres. 
A further stage of differentiation is seen in Recurvirostra 
avocetta. Dr. Mitchell has correctl}^ commented upon the spiral 
formation of the middle part of the gut, the jejunal region of the 
nomenclature adopted in the present paper. 

This bird shows the typical Limicoline characters in {a) the 
fact that the pancreas extends to the very end of the duodenal 
loop, {h) in the wide ileic loop, which is about as long as the 
duodenal, and (c) in the extent of the ileo-duodenal ligament. The 
spiral is a fairly regular one, and although certainly not longer than, 
and, I think, hardly as long as, that of the Raven, differs from it in 
the important fact that it is a permanent spiral. The several coils 
ai-e, indeed, connected together by mesentery and cannot be 
separated out without tearing this mesentery. What is a tempo- 
rary character in the more archaic forms of gut has here become a 
permanent feature. 

The Lari of Dr. Gadow's classification, which I myself prefer 
to associate more closely with the Limicolous birds, have an 
intestinal ti-act which entirely justifies the latter placing. I have 
examined Larti^s ridihundiis and L. argentatits among the Gulls. 
In L. ridihundus the pancreas, as in Limicolous birds, extends to 
the very end of the duodenal loop. The ileic loop is also wide 
and is attached for the greater part of its length by the ileo- 
duodenal ligament to rather more than the first half of the 
duodenal loop. The whole of the jejunum, which is rather long, 
lies in loose folds like the Mammalian small intestine, with no 
fixed loops at all. I could not see any trace of a spiral arrange- 
ment in this specimen. In L. argentatus, however, the jejunum 
lay in a biggish loose spiral, which was not in any way permanent. 
There is, in fact, no difference between these two species of Larus. 



78 MR. F. E. BKDDARI) OX THE 

In Tooth, it should be added, there was a distinct snpracjpcai 
kink. 

I have dissected two examples of Sterna fiuviatilis which agree 
absolutely in the characters of their gut. This genus — also like 
Larus — is very definitely like the Limicolous birds. The pancreas, 
as in those birds, extends back to the end of the duodenal loop. 
The ileic loop is wide and of about the same length as the duo- 
denal and, finally, the ileo-duodenal ligament is extensive and 
reaches nearly to the end of the ileic loop in the one case, and for 
more than halfway along the duodenum in the other. The 
jejunum lies in a rather short spiral of not more than three 
circles ; the coils of the spiral, however, are not so fixed as in 
Recurvirostra, they can be pulled apart and ari'anged in about 
three loops. This state of afl'airs has been figured by Dr. Mitchell 
for Sterna hirundo, and his figui'e would fit perfectly the con- 
ditions which I found to characterise Sterna Jluviatilis. There is 
also a supracaecal kink. 

The Auks are associated by some with the Gulls*, but by 
others they are regarded as forming a distinct assemblage t or are 
associated with the Grebes and Divers into one group Pygopodes J. 
An examination of the intestinal tract of Fratercula arctica, (text- 
fig. 26) leads me to reject the former view and to hesitate between 
one or other of the two latter classificatory schemes. At the same 
time, it must be added that there is room for divergence of opinion 
in the interpretation of certain of the loops, as will be seen from 
the following account, in which I am unable wholly to confirm 
Dr. Mitchell's account. The duodenal loop is not in any way 
remarkable and quite unfolded. The pancreas extends nearly, 
or in one specimen quite, to its end. Thereafter follow two loops, 
which belong to the middle (jejunal) region of the gut. These 
are set more or less at right angles to each other ; but the 
direction of the loop is, as I think, of less importance than the 
fact that there are two of them (and two only), which are roughly 
equal in size and very distinct. 

Moreover, these two loops are interconnected by mesenteries. 
It is obvious that we have here a close resemblance to the Grebes, 
Tachyhaptes and Podiceps §, and to those birds only among those 
whose anatomy in this i-espect is known. After these follows 
a large ileic loop, which is difi"erent in sti'ucture to that of many 
birds. Mitchell figures it as a simple wide loop. It is, however, 
long and irregularly looped, and longer than the duodenal loop. 
It appeared to me tha.t it ended in two prolonga.tions at its blind 
end, in which case there is an obvious comparison possible with 
the ileic loop in Podiceps cristatus ||, which is loosely folded at its 
blind extremity. The small intestine before the casca is thrown 
into another loop, which Dr. Mitchell has duly noted and has 

* U.ff., Mitchell, loc. cit. 

t ^.g., Beddard, 'The Stractuve and Classification of Birds': London, 1898. 

X -B. q-. Vertebrate List Zool. Soc. Lond. 1896. 

§ See" below, p. 81. || See p. 81. 



AI>IMEN"TARY TRACT OF CEUTAIX BIRDS. 



79 



irlentified with that short loop which he has termed the " supra- 
cpecal kink." In a second specimen, all these characters wei-e 
quite as plain, so that it cannot be held that the first individual 
was in any way abnormal. Probability, at any ra.te, indicates this 
conclusion. A third example, a quite young and immature bird, 
presented some slight differences which led me at first to suspect 



Text-fisf. 26. 




Intestinal tract of Fratei'cula arctica. 
Lettering as before. 

a non-identity of species. I am assured, however, that this 
suspicion is wrong. The only difierence concerned the two 
jejunal loops. Of these the first, i. e. that immediately following 
the duodenum, is much shorter than the second or more distal 
loop. The latter, instead of being a simple loop equisized 
Avith the first loop, is much longer and ha:s a kink, or sudden flexure 



80 MR. F. E. BEDDARD ON THE 

to one side, in the middle of its course. It looks, in fact, as if two 
originally distinct loops were in process of reduction to one. The 
consideration of this specimen, therefore, renders another com- 
parison possible, namely with the Cranes *, for in these birds the 
jejunum has three separa.te loops. In other respects, the third 
example of Frritercida arctica agieed entirely with the other two. 
Apart from possible resemblances to other groups of birds that 
have been indicated, the gut of Fratercida shows a feature of 
particular interest in the great length and irregular disposition of 
its ileic region. This latter is quite unattached to the duodenal 
loop, except perhaps at the very base ; it is long and lies loosely 
coiled like the Mammalian small intestine. It is, in fact, the 
loiigest section of the gut. This bird, in fact, shows an intestine 
constructed in a way which is exactly opposite to that which is 
prevalent in the class Aves. When there are tracts of primitive 
undifferentiated small intestine left it is nearly always the jejunum 
that is involved, and not the ileic region. In Fratercula the 
jejunum is specialised into fixed loops, while the ileum has 
remained unspecialised. Indeed, my experience of this structure 
among birds has only furnished one example at all parallel to the 
gut of Fratercula arctica. 

This was an example of the Pheasant, Exiplocatmis nyctlienierus. 
The Gallinaceous birds (see p. 55) ai-e very uniform in the 
disposition of their intestinal tract, and one out of two examples 
of this species which I have dissected was perfectly normal in the 
structure of the gut. A second individual, however, differed. 
The duodenal loop was immediately followed by a straight, stiffly 
fixed, descending tube of intestine, which bent back upon itself for 
a short distance and then passed into a long loosely coiled region, 
a kink became continuous ultimately with the straight portion 
of the ileum running to the junction with the cteca. This coiled 
region of the gut, although lying on the left side of the body, was 
not in any way attached to tlie duodenum. Thei'e is, therefore, 
here, it will be seen, a reversal of the conditions found generally 
among Gallinaceous birds. In this example of Fitplocamus 
nycthemerus the stiff descending intestine which immediately 
follows the duodenum obviously represents in one sense the 
equally stiff ascending piece of intestine which is in the other 
example of E. nycthemerus, and also in other Gallinaceous birds, 
parallel to and partly fixed by ligament to the duodenal loop ; 
while in the latter the loose coils which form the major part of 
the small intestine lie to the right hand. There is, in fact, a pre- 
cise reversal of the " normal " conditions. There is, as I think, an 
undoubted resemblance between this " abnormal " example of 
Eziplocamus nycthemerus and the normal arrangement of the 
intestinal tract in Fraterctda arctica. 

PoDiciPEDES. — I cannot quite explain by means of Dr. Mitchell's 
figures the intestinal loops of the two Grebes Tachyhaptes 

* Vide p. 82. 



ALIMEXTAUY TRACT OF CERTAIN BIUDS. 



81 



/lifriutili.'<* -Mid I'odicfps cn'siaiasf, of the former of wliicb s2)ecies 
I have examined two examples. The plan of the intestinal tract 
in the Grebes is very distinct and nnlike that of any other birds 
that I have studied. In Tachr/hajites the duodenal loop is con- 
siderably longer than the ileic (the precise reverse of the condition? 
obtaining in the Parrots), and the two are attached by a duodeno- 
ileic ligament which extends about halfway along the ileic loop 
(see text-fig. 27). I find only two jejunal loops (Mitchell figures 
four — two long and two short, exactly as in Ara araraibna)^ which 
are long and closely adherent for the whole of their length. The 
inner of the two loops, that which is immediately connected with 
the ileic loop, bore in one specimen a Meckel's diverticulum, as 
is indeed figured by Mitchell. The eeeca extend about halfway 
along the ileic loop, 

Text-fio-, 27. 




Intestinal tract of Tachyhaptea fluviatilis. 
Lettering as before. 



Fodicejjs cristatus conforms to the general plan seen in the last 
species, but difl'ers in detail. The ileic loop is peculiar and folded 
upon itself in a way. If unravelled it would be longer than the 
duodenal loop, and thus differs from that of Tachyhajjtes fluviatilis, 
shown in the illustration (text-fig, 27). The extent of the ligament 
uniting this with the duodenal loop is much as in Tachyhaptes. 
In the middle part of the intestine there are only two loops, as in 

* Trans. Linn. Soc. tnm. cit. p. 18tj. 
t Ibid. p. 185, «-. 0. 

Piioc. ZooL. Soc— 1911. No. VJ, 6 



82 MR. F. E. BEDDARD ON THE 

Tachyhaptes ; Dr. Mitchell figures five. Tlie two loops are wider 
than those of Tachyhaptes, but in the same way connected with 
each other up to the very end. Like Dr. Mitchell, I could find no 
Meckel's diverticulum. 

Among the Ralli there is apparently but little modification of 
the intestinal characters from genus to genus. Although I have 
not examined the actual species which Hunter has described, it is 
clear that his descriptions are in all probability perfectly correct. 
Of the " White Fulica "yPorphyrio alhus, Cuv,] " he writes * : " The 
duodenum passes down as usual, and then up, somewhat higher 
than at beginning, m.aking a sweep backwards to the loins and 
commencing jejunum. This passes down on the right side, then u^p, 
making a fold upon itself ; then a second fold, as also a third, 
which last is attached to the first fold : all these are pai-allel to 
ench other. The intestine then passes down, more in the middle 
of the abdomen, further than the former three folds, along with 
the duodenum." This fully tallies in the number of loops with 
Dr. MitchelFs statement and with my own observations ; these 
latter are, I think, worth mentioning in brief, since they further 
emphasise the uniformity of this group. 

In Hydrornis alleni, Rallus ahhotti, Povphyrio melanonotus , and 
P. madagascariensis the ileic and duodenal loops are pretty well 
the same length and are attached hj a duodeno-ileic ligament 
nearly to the end of both loops. In all of these forms there are, 
as both Hunter and Mitchell assert for species examined by 
them, three loops only in the jejunal region of the gut. The 
last of these three loops is the shortest of the three in all of 
the species which I have just mentioned. In Porphyrio (both 
species) I found a very conspicuous Meckel's diverticulum, which, 
as Mitchell correctly represents, lies near the bottom of the 
middle loop. Finally, I may remark that this middle loop, at least 
in Porphyria, is free from the other two loops (as John Hunter has 
stated), which are connected by mesentery. 

Aramides ypecaha quite a.grees with these other types and, as 
in Porphyrio, the pancreas extends to the very end of the 
duodenal loop. In this form also the first of the three jejunal 
loops is the widest, and Meckel's diverticulum occurs at about 
the middle point of the middle one of the three loo])s. 

Among the Grues I have examined among oXh&as Anthropoides 
paradisea and Balearica, which do not agree very closely in the 
nature of their intestinal convolutions with those of G7-us virgo, 
as figured by Mitchell. In both these Cranes there a,re three, and 
only three, jejunal loops, of which the first (as correctly indicated 
by Mitchell) is much the Avidest. The two following are longer 
and of equal length. On the first of these close to the blind end 
of the loop there is (in Balearica) Meckel's diverticulum. Mitchell 
figures the ileic loop as trifid. I find that in both the Cranes and in 
Grus japonicus which I have dissected (see text-fig. 18, p. 65) the 

* ' E!<says and Olisei'vations,' p. 317. 



.\l,I.Mi:.\T.\RY TIIAC'J OF CEUTAJX BIRDS. ^?> 

ileic loo|) is a double loop, the two eoin])lete vatlier narrow loops 
being not quite equisized. Anthropoides leucauchen agrees entirely 
with the other two species, and in all of them the pancreas stops 
some way in front of the end of the duodenal luop. The existence 
of three separate loops in the jejunal region is obviously a point of 
similarity between the Cranes and Rails and of difference between 
the former and certain other "Alectorides,'' e.g. Oariaraa, Bustard. 

STEfJANOPODES.- — The intestinal tract of a species of Fregata has 
been examined and reported upon by Dr. Mitchell, who, however, 
was not in a position to state precisely to wdiich species his 
observations I'efeiied. I have examined an example of Fregata 
aquila, and have something to add to the facts enumerated by 
Dr. Mitchell. He figures and refers to two cpeca. I found only one 
cfecum, which was short and sessile, upon the gut. The duodenal 
and ileic loops are simple and of a.bout the same length. The 
pancr-eas extends only about halfway down the duodenal loop 
as in Birds of Prey, and the duodeno-ileic ligament is about co- 
extensive with the pancreas. The duodenum is also connected 
with the jejunal area by a ligament which I have not found in 
other birds. 

The condition of the jejunal area is interesting when compared 
with that of other Dysporomorphous birds. It is. comparatively 
speaking, short, while that of Coi-morants and Pelicans is long. 
This comparative shortness is mentioned by Mitchell, who, however, 
has not seized upon a difference of some significance, as I think it, 
which this bird shows from its allies. 

In Phalacrocorax and Felecanus (the only other genera which I 
have examined with reference to the mattei* now under con- 
sideration) the jejunum is disposed in a consideiable series of 
closely applied regular fixed loops, as in Ducks, Storks, and some 
other birds. Fregata presents us with a stage anterior to this. 
There are no definite and regular fixed loops, but the whole 
jejunum can be disposed in an irregular circle with bulgings here 
and there. It is not a simple archaic jejunum, as in the 
Gallinaceous bird ; but neither is it the much specialised jejunum 
of other Dysporomorphee. It is hardly more advanced in the 
direction of its immediate allies than is the corresponding part of 
the intestine in an Eagle or an Owl. 

The PsiTTACi possess a complicated intestinal tract, which is, 
as I think, more correctly described by Owen than by Mitchell. 
i'or it is quite impossible to repi'esent the various loops into which 
the intestine is drawn in these birds in the fashion adopted by 
Mitchell, as will be readily seen by a comparison of the 
accompanying figure with his illustrations of Ara ararauna* and 
Stringops habroptilus f. Owen mentions the " packet of folds " 
which are alternately connected as shown in the text-figure 
appended, while Mitchell represents a series of loops sometimes 
bifurcate or tiifurcate, though stating that they are " folded 

* P. Z. S. 1896, p. 155, %. '21. 

f Trans. Linn. Sol-., Zodl. (^) viii. )i. 211, fig. '51. 



84 



MR. F. E. BEBDARD ON THE 



upon each other, and twisted and doubled in a compHcated 
fashion." It is easy to state the characteristics of the Psittacine 
alimentary tract, which in the species and genera Ara ararauna, 
A. militarise Nestor notabilis (see text-fig. 28), Chrysotis i/nornatus, 
Psephotus hcer)iatonott(,s, Callocephalon galeatum, Platycercus 
flaveolus, Cacatua sulphurea, is constructed as follows. 

The duodenal and ileic loops are single and elongated, as shown 
in Mitchell's figure, The jejunal loop consists of, pi-oximally, a 
series of three loops one within the other, of which the ascending 
limbs of two are connected respectively wdth the duodenal and 
ileic loops, and, more distally, of two sepai-ate single loops longer 
than those of the proximal bunch, which are not directly con- 
nected with each other but with the loops of the proximal packet. 
The illustration will explain these interconnections better than a 
more elaborate description, 

Text^fiff. 28. 




llitestinal tract of l^estoi" notabilis. 
Lettering as before. 

While there is in the species mentioned the general uniformity 
of structure which has just been explained, there are differences 
of small detail. Thus the two species of Ara differ, in that the 
proximal complex of loops in A . ararauna consists of only two 



ALIMENTARY TRACT OF CERTAIN BIRDS. 85 

short loops, fi-om which it follows that the two long loops inter- 
communicate directly. These facts, so far, are correctly shown in 
Dr. Mitchell's figure referred to. Conurus leucotis is precisely 
similar to Ara ararauna. 

Melopsittacus undulatus is still further reduced. The proximal 
complex is, as in the last species, reduced to two, but there is only 
one of the two distal loops left and that is shortened. 

On the other hand, Eclectus pectoralis is more complex than the 
forms already considered. Among this group of birds the liga- 
mentum ileo-duodenale is r^ather short and the pancreas extends 
to the very end of the duodenal loop, or in some cases nearly so. 
The ileic loop is often considerably longer than the duodenal. It 
seemed to me' to be' particularly long in Stringaps hdbroptilus, 
where it measured fully 8 inches in length. 

Among the Pigeons I have examined one or two forms not 
studied by Dr. Mitchell, This group contrasts, for instance, with 
the Rallidfe in the variety of intestinal patterns which it exhibits. 
This, it will be noticed, is in accord with variations in the group 
in other anatomical char'acters *• I propose, however', to deal with 
two genera in which the intestine has become shortened in relation 
to the fruit-eating habit, leaving other forms ttside until I have 
been able to make a moi^e comprehensive study. In one of these, 
Ptilopus hellus, the reduction in length, coupled with great increase 
in calibre, has not go'ne so far as in the genus Carpophdga, with 
which I shall deal later. In Ftilopiis belhcs the duodenal loop is 
quite well mai-ked, though short and rather wide' ; the pancreas 
extends to its very end. The jejunal region which follo\vs is also 
distinct from it ftnd fro'm the ensuing ileic loop. The jejunal 
region consists of a single loop only, the' two limbs of which lie in 
close apposition, but can be considerably separated without tearing 
any membranes. This region of the gut is therefore, in consisting 
of a single loop without further complications, precisely like that 
of the Bustards and Cariamas. Tlie ileic loop is also well marked 
and rather wide ; it is much longer than the duodenal. The ileo- 
duodenal ligament is present, but not very e'xtensive. There is 
also a ligament uniting the jejunal loop' to the entering limb of 
the ileic loop for about halfway down the' latter. The plan of 
the intestinal tract in this bird is therefore a i-athef primitive one, 
reminding us of that of Otis and Gliunga and of the Ficopass6res. 
In fact, the degeneration of the intestine has resulted in the throw- 
back to a presumsibly earliei- state of afFairsv 

The genus Carpojuhaga shows a further degerieration of the 
intestinal tract, which is well known to fee very short in 
this genus of Pigeons f. The shortening is allso accompanied by 
widening. 

I have examined CJ. ceuea and C. concinmcc- In both of these 
the duodenal loop has vanished and is represented perhaps by the 

* Gan-od, " On some Points in the Anatomy of the Columhte," P'. Z. S. 1874, p. 249. 
t Cf., c. (/., GarroU, "Notes oU tht Oizzaid &*, of Caryoj/hrt^a latrans," P. Z. iS. 
1878, "p: lOk 



86 MR. F. E. BEDDARD OX THE 

.slightest kink in the ahmentary tube. The rest of tlie tube is 
disposed in four parallel and transversely arranged lines, which on 
further examination are seen to form a. spiral and which end in 
a longitudinal section of gut running to the cloaca. I cannot 
distinguish in this a jejunal from an ileic region. It is interesting 
to compare with this " degeneration " of the intestinal tract in 
the fruit-eating Pigeons the coiresponding alterations met with 
in the fruit-eating Passerine yEluro&dus. In that biixl there is 
a vestige left of the duodenal loop, to the extremity of which the 
pancreas reaches. Indeed, the rest of the gut has retained, 
though in a reduced condition, the two recognisable divisions, of 
which the ileum is represented, as in the primitive Picopasseres, 
by f\ straight tract parallel with and connected by the usual 
membrane to the duodenal loop. 

§ Some General Considerations. 

The new facts which have been described in the foregoing 
pages give rise to certain reflections upon the affinities which 
they appear to indicate between different families of birds and 
upon the course pursued in the evolution of the gut of birds. 
The facts, so far as they are known, do not appear to me to lead 
to the establishment of a phylogenetic scheme, even of the 
evolution of the gut only, so elaborate in the setting forth of 
details as that which is drawn up by Dr. Mitchell as the result of 
his own labours in this department of anatomy. ISTevei'theless, it 
does seem possible to indicate certain stages in the evolution of 
the intestine, and here and there are indications, already to some 
extent considered, of affinities between different Avian families. 

§ The Primitive Form of the Intestine in Birds. 

It is obviously necessary, before considering the features which 
are the most primitive in the Avian alimentaiy tract, to get a 
clear notion of the essential differences which distinguish the 
alimentary ti-act of Reptiles from that of Birds ; by this means we 
shall evidently arrive at the essential resemblances. Dr. Mitchell 
distinguishes the bird's intestine thus: — " It is distinguished from 
the intestinal tract of reptiles chiefly by the fact that the three 
divisions — the duodenum, Meckel's tract, and the rectum — are 
sharply marked off" one from the other." These lines are written 
of Palamedea, which that author regards "as representing closely 
the ancestral type." It appears to me, however, that while 
Palamedea is undoubtedly an ancient type, the definition used by 
Dr. Mitchell is not a correct one. For, while in, at any rate, the 
majority of Lizards known to me there is a very marked dis- 
tinction between the small intestine and the large, the Crocodiles 
show a further differentiation ; for they show a, very well-marked 
duodenal loop as well. The divisions of the alimentary tract 
therefore do not enable us to distinguish bet\veen Birds and 



ALl.MKXTAKV TRACT OK CEKTAIX lilltOS. 87 

Reptiles. Indeed, in an earlier paper*, Dr. Mitchell lias himself 
sketched in a perfectly correct fashion the aliraentaiy tract from 
an Alligator, illustrating the facts to which I have just directed 
attention. Fi'om a gut like this it appears to me to be only just 
possible to distinguish that of Gasuccrivs t in its general charac- 
teristics, among which I do not include the well-developed cjsca. 
The only difference that I can detect is a closer approximation 
between the ileic and duodenal regions in Casuarkis, already, 
however, mai'ked, though to a less extent, in the Crocodilidae, 
which, of course, foreshadows the very close association found in 
all other bii-ds. This association, caused by the outgrowth of the 
long middle part of the small intestine from a short region of 
the primitively straight gut, naturally bi-ings about the com- 
mencement of the formation of the fixed ileic loop, so conspicuous 
a character of the alimentary tract of other birds. It is most 
interesting to notice that among Crocodiles there is, in some 
species at least, a quite distinct ileic loop, related perhaps to this 
same association between the ileic and duodenal regions, which 
is not, however, as has been already remarked, so close among 
Birds. It is to be noted that here as elsewhere the closest asso- 
ciation of Birds and Reptiles is shown, thoroughly justifying the 
views of Cope, Huxley, and others. It may be admitted, therefore, 
that Casuarius is, at any rate, one of those birds whose intestinal 
tract, both arrangement and convolutions, hardly differs from 
that of Reptiles, and is therefore primitive as compared with that 
of many other birds. Nor, indeed, is there so far any very great 
difference from the most primitive form of the gut in Mammals, 
where, as in Casuar^iibs and Crocodilus, the entire intestinal tract 
is borne upon a continuous mesentery. 

§ The Course of the Evolution of the Gut. 

From the simple conditions which obtain in Gasioarius the 
more complicated intestinal tract of other birds can be derived : 
and an almost complete chain of intermediate stages is exhibited, 
even among the few genera which I have had the opportunity of 
studying. It is from this point that the characters of the intes- 
tinal tract in Birds diverge from those of Mammals, the Reptilian 
conditions being left behind by both groups of Vertebrates. It 
may be convenient at this stage to point out the essential 
differences which distinguish the intestinal tract of Birds from 
that of Mammals. It has already been pointed outi that one 
difference is to be seen in the fact that among Mammals the 
permanent loops of the large intestine distinguish that gut from 
the small intestine, where there are no such permanent loops ; 
whereas in Birds it is the small intestine only which ex- 
hibits these permanent loops. These characters, however, though 

* P. Z. S. 1896, p. 137, fig. 1. 

t I have cvamined the species C. rotJixr.hiJdi, C. intensus, and C. wesfcrmaitni. 

X E-J; Gcgenbaur, Vergl. Auat. d. Wirbelth. 



CO MR. F. E. BEDDARU OX THE 

distinctive in the negative sense are not universal. There are 
whole orders of Mammals, e. g. Oarnivora, Marsupials, Primates, 
Edentata, in which the colon has no permanent loops, while in 
many birds, e. g. Gallinaceous birds, certain Picopasseres, the 
small intestine has no fixed loops. In addition to this very 
important difference, there appear to me to be two other distin- 
g-uishing featvires in the alimentary systems of the two orders, 
which are perhaps equally important and are certainly more 
regularly met with. In all Mammals above those few forms 
(e. g. certain carnivorous Marsupials, certain Edentata * ), which 
have so far retained the Reptilian characters of gut, the whole 
intestine is rotated upon itself and the rotation is fixed and the 
change of position of the various sections of the gut retained by 
the ligamentum cavo-duodenale which moors the end of the 
duodenal loop to the colon, mesocolon, or median dorsal body-wall 
on or near the postcaval vein. This ligament correlated with 
the said rotation is universal among the higher and present in 
most of the lower Mammals, In Birds, on the contrary, there is 
no such tract of mesentery fixing the duodenum to the colon. So 
far, in fact, the bird's intestine has retained the primitive Reptilian 
condition. The bird's intestine, however, usually has what the 
mammal's intestine has not, a duodeno-ileie ligament. This doubt- 
less is the ijhysiologieal equivalent of the duodeno-colic ligament 
(as Mitchell t has pointed out), in so far as it serves to anchor the 
perhaips otherwise inconveniently long and coiled small intestine. 
It may also perhaps be argued from this that a short intestine 
(<?. g, many Picopasseres) is shown to be a secondary state of affairs, 
from the very fact that it is in those birds provided with a duodeno- 
ileic ligament, which may not be a mechanical necessity. For 
the existence of the ligament in question may be due to mecha- 
nical needs in an ancestor with a long small intestine. In any 
case, the morphological fact is to be noted and it constitutes a real 
difference between the Mammalian and Avian gut. 

In very nearly all birds whose intestinal tract is at or above 
the level of that of the Cassowary, the end of the small intestine + 
is attached by a mesentery of vaiying degree of development to 
the duodenum. This, without any further specialisation, is the first 
stage in the evolution of the gut from its simple archaic form. 
This stage characterises the hypothetical Gallinaceous birds in 
which it is universally present and quite similar in alL 

This simple stage, which we may term Stage A, is also found in 
other groups, but it does not occur in all the members of a.ny 
other group as it does in the case of the Gallinaceous birds. 
Among the Struthious birds, for example, we have it in the 
Cassowaries, Ostrich, and Apteryx §, whose intestinal tracts are 

* For a general sui-vey, see Klaiatsclij Morpli, Jabrb.- xviii. 1892, and myself in 
P. Z. S. 1908, p. 568 &G, 

t Trims. Z. S, xvii. p, 524, 

[J; The chief exception known tO' me is fnniislied by Fratercula arctica, the 
remarkable characters of tlie intestine of which bird I have already coanmentcd 
upon {supra, p. 78). 

§ The condition of Rhea requires perhaps further study. 



ALIMENTARY TRACT OF OERTAIX BIRDS. 89 

precisely similar so far to that of the Gallinaceous birds. We 
have in certain Picopasseres (e. g., Melanerjyes) the same state of 
aft'airs, coupled in this case with a considerable shortening of the 
gut. Among the Limicolae the genus Pltcvianus is also in this 
stage. 

The next stage, which may be called Stage B, is like the last, 
save for the fact that the ileic loop is definitely formed. The 
jejunal region remains unspecialised. To this stage, we refer the 
gut in the majority of Picopasseres, including the Cuckoos and 
Plantain -eaters. The "Alectorides" (in the sense in which I ventui^e 
to use that term in the present communica,tion) seem, but perhaps 
only seem, to belong to this stage. Among the Limicolous birds 
iSa?'ciophoi-}ts find Gulls appear to belong here. Perhaps we should 
also place in this assemblage the Dyspor-omorph Fregata. 

Stage C is a slight advance upon the foregoing. It is exem- 
plified in certain Owls and Hawks, where the ileic loop is fully 
difi"erentiated off and attached in the usual way to the duodenum, 
and where the jejunum is largely laxly coiled without any definite 
loops, save a single loop, and that not a very well-marked one, 
which occui-s at the commencement of the jejunum. I have not 
noticed this kind of intestine in any other group, except the 
Nocturnal and Diurnal Birds of Prey. 

Stage D might possibly be further divided up, but for the 
present I do not see a clear Way through the great variations 
which the intestine of the more complicated forms shows. In all 
of them the jejunum has become differentiated into fixed loops, 
which vary in number, in relative length, and in their relations 
one to the other. The majority of the larger birds belong to this 
stage, as, for instance, the Cranes, Rails, Ducks, and Storks. 

§ TJte Mutual Affinities of Avian Families jicctged hy the 
hitestinal Coiivohitiovs^ 

The known facts do not, a«s I think, permit of any complete 
scheme of classification of Birds by means of the variations in 
the coils of the intestinal tract. Here and there, however, there 
would seem to be such indications, which are tolerably well 
marked. More frequently, however, either a general plan runs 
through a seiies of two or three groups, which makes any 
definite placing of these groups in reference to each other 
difiicult, oi" a complete isolation is shown. The most salient 
instance of the latter conclusion is undOTibtedlj' the group of 
Parrots, whose intestinal coils are constructed upon a. plan which 
is apparently universal in that group, but totally unlike anything 
which i&' to be found in any other group. The afiinities of the 
Psittaci have been very variously interpreted *, but it is clear 
that the gut does not enable one to decide upon any of these 
divei'se views.- I cannot at all agree with Dr. Mitchell in saying 

* Miuij- or most of tlicse ojiiiiions arc mentioned by Fiirlirin<rcr in liis 
monumeiita'l' worlc. ' Uii'tursuehuimcn neber Morpli. ii. ^yst. dev Vogcl,' Amsterdam, 
1888. 



90 MR. P. E. BEDDARD OX THE 

that " the relation to the common type is, however, easily made 
out " *. 

In the same way, the Ralli are a quite circumscribed group 
judged by their intestinal coils, which bear only a general resem- 
blance to other groups and, indeed, to no group in particular. 
Their characters are constant throughout the group, so far as 
known facts enable us to make a statement. The most that can 
be said is, perhaps, that they are nearer to the Grebes than the 
latter are to any other subdivision of the Bird tribe. On the 
other hand, it seems to me to be quite clear that if judged by 
their intestinal coils the newer ideas with respect to the New 
World Vultures and other Accipitresf must be abandoned, as 
Dr. Mitchell has correctly pointed out ; it is necessary to revert 
to the older view which regarded the Condors as merely Viiltures. 
In the same way, although in this Dr. Mitchell does not agree 
with me, the older opinion as to the Owls, that which placed 
them close to the Accipitres and not in the neighbourhood of 
vai-ious Picarian genera, is most certainly justified by the close 
similarities in the mode of arrangement of the intestinal loops. 
At the same time, it is also easy to distinguish these two groups 
by the small but constant characters afforded by the ileo-duodenal 
ligament. 

And, again, it is by no means possible to distinguish by cha- 
racters that carry any conviction the intestinal tract of a Grebe or 
Tern from that of the Owls on the one hand or large Passerine 
birds on the other ; while the Gulls and Terns on their side 
offer resemblances to what I venture to term " the other " Limico- 
line birds. Negative features are perhaps more salient in making 
a brief sui-vey like the present. Thus it is clear that the 
Tinamous are quite unlike the Gallinaceous birds and that the 
genus Turnix is equally to be removed from that assemblage. 
The Bustards and Cariama, moreover, show no. particular likeness 
to the Cranes, though the first two seem to be closely allied to each 
other, as I have already pointed out. It is noteworthy that all the 
four types J of Struthious birds differ from each other as much 
as would seem to be possible in view of the undoubtedly primitive 
characters of the gut in all of them, with the possible exception of 
Rhea. 

§ The Relationship between the Gut and the 
Nature of the Food. 

"When we contrast the intestine of a Penguin with its enormous 
series of closely adpressed straight loops and the intestine of a 
Bustard with only three short intestinal loops, there would appear 
to be a very marked difference between a fish-eater and an 
omnivorous bird, and thus a close relationship between the form 
and length of the gut and the nature of the food eaten by its 

* P. Z. S. 1896, p. 155. 

t These opinions are so well known that I need not quote what would have to be 
a lonp: list of books and memoirs. 
X Casuarius and Droitiicus belong, of coursBj to the same type, 



ALTMIOXTARV TRAf'l OF CKRTAIX BIRDS. 91 

possessor. There is not, liowever, in .ill cases a relationship 
which is plainly to be recognised between differences in diet and 
differences in the intestinal part of the alimentary tract, and on 
the other hand a totally different diet sometimes coincides with a 
close similarity in the intestinal convolutions. Phylogenetic 
I'elationship appears to me to have much more to do with these 
similai-ities and differences in the gut. Furthermore, the way in 
which the intestinal tract is modified in accordance with the diet, 
when it does .appear to be so modified, seems to have pursued a 
<liSerent path in different groups in some cases. Of fi.sh-eating 
birds, for instance, the general idea is that the gut is long, and 
that undoubtedly is the case with the Penguin and the Cormorant- 
Pelican group. But then in these groups no representatives a,re 
known which are not fish-eaters. Directly we come to the 
consideration of groups of birds which contain fish-eating genera 
and genera whose food is not fish, we are sometimes met by a 
totally diflerent state of afiairs. 

According to the statistics collected by Mr. Newstead*, the 
Terns (of three species) a,re exclusively fish-eaters. Yet their gut 
is not markedly and indeed hardly at ail different from that 
of the Avocet, which devours aquatic insects, and some other 
Limicolfe which select a similar diet. The .Kingfisher {Alcedo 
ispida), which is, according to the same authority, practically 
entirely a fish-eater, has a gut which is very like that of the 
omnivorous Corvines, and has, moreover, a much shorter spiral 
jejunum than in the differently feeding Halcyon sancta. The 
Toucans t are mainly fruit-eaters, though, like so many birds, 
they will vary this diet with animal food. And yet their intes- 
tinal tract differs very little from that of Podargus, which is 
presumably not at all a fruit-eater, but subsists entirely upon 
insects and other animals. Again, the Touracous X are fruit- 
eating birds ; but their gxit is like that of a vast series of Pico- 
passerine birds which feed upon all kinds of food. 

In asserting that the gut is short in all purely frugivorous and 
insectivoi'ous birds. Dr. Gadow practically admits how little stress 
can be laid upon the relationship between length of gut and the 
nature of the food. For the nature of the diet in each case is as 
different as possible. Nor can any general principles be stated as 
to the complication of the gut in families of birds which live 
differently. Thus the plan of the gut in Apteryx is practically 
identical with that of the Gallinaceous birds, and the character 
of the food differs. On the other hand, the pattern of the gut in 
Accipitrine birds is not dissimilar to that of Owls, and here w^e 
have a general similarity in diet. It is, in fact, not possible to lay 
down general rules which have not copious exceptions. Many of 
these exceptions can be gathered fi'om the foregoing pages. 

* Supplement to th(> .Journal of the noavd of A.cjricultuve, vol. xv. No. 9 (1908). 
t Newton, ' A Diftion;ivy of Birds ' (Loudon, 1893), sub voce ■•Toucan." 
J; Id. thirl. ^ " Tounicou." 
■ § Id. ibid., " Digestive .System.' 



92 ON THE ALIMENTARY TRACT OF CERTAIN BIRDS. 

§ Summary of Facts relating to the Intestinal 
Coils of Birds. 

We are in a position, I think, to lay down with confidence the 
following genei-al statements with regai-d to the intestinal tract 
of the Class Aves. These statements are deduced from the 
memoirs of others who have written upon this subject, as well 
as from the facts ascertained by myself, and embody in a brief 
form what is known upon the subject dealt with in the present 
communication : — 

(1) There are no essential difierences between the intestinal 
tract in Birds and in Crocodilia. The most complicated alimen- 
tary tract in Birds can be derived through a series of stages from 
the simple Crocodilian form. 

(2) The intestinal tract of Birds diffei's from that of Mammals 
in that there is never in the former, as there is generally in the 
latter, a rotation of the gut coupled with an attachment of the 
duodenum to the colon or mesocolon by a duodenal caval ligament. 
On the other hand, there is nearly alwa^ys in Birds an ileo- 
duodenal ligament which serves to hold together the gut and 
which is wanting in Mammals. Furthermore, in Mammals the 
colon may be specialised into fixed loops, while in Birds such 
fixed loops are found only in the small intestine. 

(3) The small intestine of Birds can, with rare exceptions, be 
distinguished into duodenal, jejunal, and ileic regions. 

(4) The duodenal region consists of a long well-defined loop 
(only absent in certain fruit-eating Pigeons), which is usually 
straight, but sometimes (e. g. Milvus) shows indications of — or a 
pronounced — spiral arrangement, as it does also in certain 
Crocodilia,. It is wider or narrower in different groups, and the 
enclosed pancreas extends a greater or a less way towards the 
extremity of the loop in different birds. 

(5) The jejunal region is sometimes (e. g. Gallinaceous birds) 
marked ofi" from the duodenal by a sudden diminiition in calibre. 
It is usually the longest section of the gut ; but is sometimes as 
short or even shorter than either the duodenal ot ileic region or 
both. It may be a tract of intestine without any fixed loops (as 
in the Gallinaceous birds, various Passerines, &c.), but if of con- 
siderable length is usually specialised into a series of two or more 
fixed narrow loops, which may be interconnected by secondary 
mesenteries in different ways, distinctive of difierent groups of 
birds. An intermediate condition is seen (e. g. among Accipitres 
and in Fregatct), where the jejunum is not arranged in lax and 
alterable coils as iii the Gallinaceous birds, but is stiffened here 
and there into wide loops, which are not so sharply marked off 
as in the more specialised birds and are not interconnected by 
secondary mesenteries. 

(6) The ileic region is not always quite distinct from the 
jejunal, and degrees of distinctness occur. In less specialised 
intestines there is no definite loop, but the end of the ileum is 



ON SPOTTED HY.ENAS IN THE BRITISH MUSEUM. l)?{ 

attached for a variable distance by a secondary mesentery to tlie 
duodenal loop. In more specialised intestines the ileic region 
forms a distinct loop which is narrower or wider, and which is 
equal to, longer, or shorter than the duodenal loop, to which it is 
attached for a greater or less length by the ileo-duodenal ligament 
already mentioned. Not infrequently the ileic region consists of 
two loops, of which one is often small and lies just above the 
ca?ca. 

(7) The plan of the gut is constant (except for very slight 
variations) in all the members of certain groups. This is the case 
with the Psittaci, Galli, Raptores, Striges. In the case of other 
groups, there are considerable divergences of structure within the 
group : this is the case with the Picopasseres, Limicolpe, Grues, 
Struthiones (if, that is to say, the two latter groups can be 
regarded as natural groups, which is open to doubt on other 
grounds). 

(8) A comparison of the intestine of Birds Avith that of 
Reptilia (especially Crocodilia) allows of the recognition of more 
and of less primitive types of intestine. The most primitive type 
is found in the Cassowaries, Strutkio, Apteryx, and all the 
Gallinaceous birds ; and is also seen among the Picopasseres 
(Woodpeckers) and Limicolse {Pluviamts) . Most Picopasseres 
and the Birds of Prey (nocturnal as well as diurnal) show a rather 
more specialised form of intestine. In the remaining groups of 
Birds the intestine is more specialised still and in several different 
directions. 

(9) Certain classificatory results seem to follow from a 
comparison of the differences exhibited by the intestinal tract. 
Thus, the resemblance of both Cuculi and Musophagi to the 
Picopasseres, and the likeness between all the Accipitres (New 
"World and Old World, nocturnal and diui'iial) are remarkable. 
The close likeness between the Bustards and the Caiiamidse is 
to be commented upon. The Passerine character of the gut of 
Turnix and the possible likeness between Cryjyturus and Rhea 
seem also to be shown. 



7. On the Specimens of Spotted Hyasnas in the British 
Museum (Natural History). By Prof. Angel Cabrera, 
C.M.Z.S. 

[Received November 5, 1910: Read November 29, 1910.J 

Every zoologist working on the Spotted Hysenas with suitable 
material from different localities feels the convenience of re- 
cognising several local forms. As early as in 1812, two of them 
were admitted by Cuvier, and in modern times no less than nine 
other " species " have been described. It is not easy to say with 
which form Erxleben's Hycena crocuta, afterwards the typeof the 
genus, or subgenus, Crocaia, must be identified, as the species was 



94 PROF. A. CABKEEA ON 

based on the " Spotted Hy«3na"of Pennant *, and this author 
gave no definite locality, saying only that the animal is fovmd in 
" Guinea, Ethiopia, and the Cape." That indication practically 
embraces all the African countries known in Pennant's time, 
excepting only Barbary and Egypt. As to the original de- 
scription, made from a specimen shown in London some years 
before, it runs thus : — 

" Short black mane : hair on the body short and smooth : ears 
short and a little pointed ; their outside black, inside cinereous : 
face, and upper part of the head, black : body and limbs reddish 
brown, marked with distinct round black spots ; the hind legs 
with transverse black bars ; tail short, black, and full of hair." 

Now, I have never seen, nor found described, a Spotted Hysena 
with black mane. Young specimens commonly have dai-k hairs 
in it, producing a general blackish tinge, but it seems clear that 
Pennant's specimen was not young, as in the description it is 
afterwards stated that it was bigger than the striped species, and 
the author says about the latter in a previous page that it is 
larger than a big dog. It is therefore necessary to suppose either 
that Pennant spoke from memory and forgot some details of the 
coloration, or that the actual specimen represented a form quite 
unknown to modern naturalists. The latter view being a very 
unlikely one, I prefer to think that the desci'iption was written 
from memory only, under the impression of a reddish-brown 
animal spotted with black, and perhaps a not quite developed 
specimen with a little of the juvenile dark hair in the mane. 

This determination being adopted, it seems to me very probable 
that the specimen alluded to came from Senegambia, although 
this locality is not mentioned among those given by Pennant as 
inhabited by Spotted Hytenas. Senegambia and the Cape were, 
during the eighteenth century, the two countries that chiefly and 
almost exclusively furnished the European menageries with 
African animals. But in the Cape Hytena the ground-colour 
is a dirty yellowish which nobody would call reddish brown, 
whereas this rather indefinite designation may be correctly 
applied to the peculiar colour, intermediate between dark 
cinnamon and raw umber, of the Senegambian Hysena. The fact 
that Pennant did not include Senegambia in the habitat of the 
species is of little, if any, impoi-tance, as he compiled the geo- 
graphical distribution from the works of Bosman, Kolbe, &c., and 
Avas evidently unaware of the provenance of the specimen he saw 
in London. In his ' Game Animals of Africa,' Mr. Lydekker says 
that the typical Hycena crocuta is the form found fi'om Southern 
Egypt, across Central Africa, to Senegal in the west and the 
Transvaal in the south. I cannot agi-ee entirely with such a 
conclusion. As will be seen below. Spotted Hyaenas fi'om the 
Nile Basin and East Centiul Afi-ica ai-e verj' different in coloui- 
from the animal described by Pennant and Erxleben, and therefore 

* ' History of Qiuulrupcds; i. (1781) p. 252. 



SPOTTED JIY.EXAS IX THE BRITISH MUSEUM. 95 

these countries at least must be discarded. The supposed dis- 
tribution being restricted in this manner to Central Africa and 
Senegambia, it becomes clear that, as Centi-al Africa was totally 
unknown to Europeans in Pennant's time, the only conclusion at 
which we can arrive from Lydekker's statement is that which I 
have adopted. Crocntta crocuta may therefore be selected as the 
name for the Senegambian Spotted Hyajna, at least until a 
stronger argument can prove that this view is not the right one. 

As I have said above, Ouvier, in his ' Ossements Fossiles,' 
distinguished two different forms of Spotted Hytenas, a grey 
one and a reddish one. In the second edition of the same work, 
he states that his reddish Hytena is frequently found about 
the Cape, but the source of this knowledge is not given. The 
first exact description of the Cape Hysena is by Desmarest*, who 
named it ffycena cajyensis. As to H. rvfa, of the same author, 
based on Cuvier's i-eddish Hysena, its exact locality being unknown, 
it is best to consider it as a synonym of true crocuta. Boitard t 
gives the three names to one and the same animal, and describes 
a yellowish Hyaena from the Cape as H. citvieri, this name 
becoming thus a synonym of Desmarest's capensis. 

Since the publication of all these old names, no other splitting 
of the group has been attempted till 1900, when MatschieJ de- 
scribed five so-called new species : Crocuta wissmanni, from German 
"West Africa ; C. gariejjensis, from the Orange River ; C ger- 
minans, from Gei'man East Africa ; and (7. thierryi and 
C. togoensis, from Togo. In a subsequent paper §, the same 
zoologist named the form from Kamerun C. noltei, Satunin || has 
given the name leontiewi to the Abyssinian Spotted Hyaena,, a,nd 
Lonnberg ^ has described two other forms from East Africa, 
C. kibonotevisis, from Kibonoto plains, and C. j^anganensis, from 
the Pangani River. 

It is impossible to decide now, without the comparative study 
of large series of specimens, and especially of skidls, whether all 
these forms are true different species, or whether they are local 
races of one or of several species. From the material in the Bi-itish 
Museum, which I have been able to examine through the kindness 
of Mr. Oldfield Thomas, F.R.S., it appears that two different types 
of skulls may be distinguished : — a broad one, in which the width 
of the palate across the upper carnassials is practically equal to or. 
a little greater than the length of the lower tooth-series exclusive 
of incisors ; and a narrow one, in which the width of the palate is 
equal to or a little less than the length of the upper tooth-series. 
This latter always being 4-15 mm. shorter than the lower tooth- 
row, it is clear that the difference between the two types can be 

* ' Mammaloo;ie,' i. (1820) p. 216. 
t Le Jardin des Plaiites, 1845, p. 232. 
X SB. Gesellsch. Nat. Fr. Berl. 1900, pp. 18-58. 
§ L. c. 1900, p. 211. 

i| ' Zoologischcr An/.ei.srer,' xxix. (1905) p. 556. 
% Sjosted't, Kilimanj. Mcru Expcd. 1908, pp. IB-lS,. plf^. 5 & 7. 



96 PROF. A. CABRERA OX 

detected at first glance. Moreover, the ratio between the zygo- 
matic breadth and the condylo-basal lengtli is 65 to 72'5: 100 in 
narrow skulls, and 70 to 75 : 100 in broad skulls. Of course, I 
refer solely to adult specimens ; in young skvills the proportions 
are very changeable, according to the age. 

Three only among the eleven forms described up till now 
seem to be represented in the British Museum collection, as 
follow : — • 

Crocuta capensis Desm. 

Two adult specimens, one from the Cape (B.M. no. 46.8.3.3, 
Tamer), and another from the Pongola River, Zululand (B.M. 
no. 2.2.8.1, D. Bruce), and a young one from the Cape (B.M. 
no. 37.9,26.90, Turner). Gi'ound-colour dirty cream-bufl"; spots 
small, round, numerous, and of a pale hair-brown ; mane dirty 
ochre-yellow ; snout and feet dai-k-coloured, between sepia and 
hair-brown. Skull of the broad type. 

Skull-measitremenis of no. 46.8.3.3. — Condylo-basal length, 
236 mm.; zygomatic breadth, 176; interorbital breadth, 57; 
postorbital constriction, 42 ; rostral breadth on canines, 64 ; width 
of palate across the carnassials, 114; mandible, fi-om condyle, 
180; upper tooth-series*, 100; lower tooth-series. 111; upper 
carnassial, 35 x 22 ; lower carnassial, 30 x 12. 

A specimen from the Cape in the Madiid Museum of Natural 
Science is entirely like the British Museum specimens. The 
figure given by F. Cuvier in his ' Histoire Natui-elle des Mammi- 
f^res,' after a living animal obtained in the saine locality by 
Capt. Baudin, is not good, the difierence in height between the 
fore and the hind quarters being too exaggerated. 

Crocuta wissmanni Matsch. 

An old specimen from Linyanti (B.M. no. 0.10.3.1, P. C. Eeid). 
Ground-colour ochraceous ; under surface, from the throat, pale 
cream-colour. The spots are very dark brown and show a 
tendency to form longitudinal rows on the flanks. On the limbs 
they are darker, almost black, and reach a lower level than usual 
in the group. Mane ochraceous. Snout dark brown. Feet buff- 
yellow. Skull of the narrow type. 

Skull-measurements. — Condylo-basal length, 256 mm. ; zygo- 
matic breadth, 185; interorbital breadth, 64; postorbital 
constriction, 53 ; rostral breadth on canines, 66 ; width of pala,te 
across carnassials, 112; mandible, from condyle, 1 90 ; upper tooth- 
series, 113; lower tooth-series, 117; upper carnassial, 35 x 20 ; 
lower carnassial, 32 x 11. 

It is not without hesitation that I call this specimen 4OTssmaw?w, 
as it lacks the black feet which Matschie says are characteristic 
of the German West Africa Hysena, and which are to be seen in 

* In dcscvihing Carnivora, I always measure the tootli-rows from the front of 
the canine to the back part of the hindmost cheek-tooth. 



SPOTTED HY.EXAS IX THK BRITISH MUSEUM. 97 

Schreber's plate xcvi. (Saugtli. ), with which toissmanni is iJentiiied 
But the ochraceous ground-colour, the paleness of the under parts, 
and, moreover, the relative proximity of Linyanti to Epukiro, the 
type-locality of toissmanni, prevent me from giving it a new name, 
at least until some other specimens are available. Bv the way, 
the material on which Crocuta loissmanrd was based is far from 
good. The type is a skin without skull, and Prof, Matschie is 
not sure that the skull from Windhoek, described in his paper, 
really belongs to the same form. 

CrOCCTA LICOXTIKWI iSiituniu. 

A skull, without skin, from Abyssinia (B.M, no, G9.2.2.13, 
Jesse). Its dimensions answer rather well to those given bv 
Satunin for his specimen 5784. It belongs to the narrow type 
and is broken behind, it being, therefore, impossible to measure 
its condylo-basal length. 

Skull-measurements. — Zygomatic breadth, 153 mm.; intei'orbital 
breadth, 55*5 ; postorbital constriction, 41 ; ro.stral breadth on 
canines, 58 ; width of palate across the carnassials, 99 ; mandible, 
from condyle, 172 ; upper tooth-series, 98 ; lower tooth-series, 103 ; 
vipper carnassial, 36 X 19 ; lowei- carnassial, 26 x 10. 

Another skull, without skin, fi'om the White Nile (B.M. no. 
2.8.5.4, Maj. Dunn), seems to belong to the same species ; whereas 
another from Bar-el-Zaraf (B.M. no. 0.8.6.2, Stanley Flotoer) 
evidently represents a different form, perhaps the Huflanese 
Hyeena, which I do not dare to name without knowing the colours. 
It is of the broad type, and undoubtedly came from a very big- 
animal. 

There are in the British Museum, besides the specimens just 
mentioned, some otliers representing three different and ap- 
parently new forms. In describing them, it is oiily provisionally 
that I use binomial names. 

Crocuta rufopicta, sp. n. 

Diagnosis. — A very pale reddish form with red spots, quite 
different from the grey Hytenas of Abyssinia and Uganda. 

Colour. — Ground-colour pale ochraceous buff, verging to tawnv 
ochraceous on the back and the mane, and fading to pale buff on 
the under parts. Spots small, very scattered, and ochraceous 
rufous in colour. Feet ochraceous. Tip of the tail blackish, as 
usual in Spotted Hysenas. 

Skull. — Palate nari-ow, its gi'eatest breadth being rather less 
than the length of the upper tooth-series. Bullfe elongate ; 
paroccipital processes considerably drawn backwards, their hind 
outlines appearing, when the skull is I'esting on its basis, aa 
oblique lines. 

Skull-measurements (of type). — Condylo-basal length, 240 mm. ; 
zygomatic breadth, 159 ; interorbital breadth, 53'5 ; postorbital 
constriction, 40 ; rostral breadth, 59 ; width of palate across the 
carnassials, 97; mandible, from condyle, 178; upper tooth scries, 

Proc. Zool. Soc— 1911. No. VIl/ 7 



98 ■ PROF. A. CABRERA ON 

103; lower tooth-series, 106-5; upper car nassial, 36x20; lower 
carnassial, 28 X 11. 

Hah. Odixeina, Boran Country. 

Tyjie. Old female. B.M. no. 9.6.1.14. Collected by Mr. Drake- 
Brockman. 

Remarks. — The type-skin lacks the head and fore part of the 
neck, but it is so different from all the other Spotted Hyaenas 
that, although the only specimen and an incomplete one, it 
deserves to be considered a new form. C. kibonotensis, pale 
reddish in colour like ?^nfopicta, has the spots dark brown, not red, 
and its skull, an excellent photograph of which has been published 
by Lonnberg, is also different from that of the present species or 
race. 

Crocuta thomasi, sp. n. 

Diagnosis. — A pale grey, black-spotted Hysena, with the skull 
of the narrow type. 

Colour. — General colour pale gi'ey, strongly suffused with buff" 
on the fore quarters, and marked with large black spots, very 
irregular in form, some of them being elongate and even some- 
what linear. Mane dirty ochre-yellow, fading to cream-buff" on 
the sides of the neck, where there are two irregula.r rows of faint 
burnt-umber spots. Feet and legs to near the elbow and the 
knee dark clove-brown, almost black. Tail pale grey, blotched 
with small blackish spots, and with the distal half black. 

Skull. — Palate narrow, but not so much as in C. ru/ojncta, its 
largest breadth being practically equal to the length of the upper 
tooth-row, and even considerably exceeding it in young specimens, 
in which the carnassial is not yet quite developed. Hinder out- 
line of paroccipital processes forming a vertical line. 

Skull-measureinents (of type). — Condylo-basal length, 245 mm. ; 
zygomatic breadth, 165 ; interorbital breadth, 54 ; postorbital con- 
striction, 41 ; rostral breadth on canines, 58 ; width of palate 
across the carnassials, 101 ; mandible, from condyle, 175; upper 
tooth-series. 100; lower tooth-series, 107; upper carnassial. 
35 X 18 ; lower carnassial, 26 x 10" 5, 
Hah. Ankole, Uganda. 

Type. Adult male, B.M. no. 1.8.9.27. Collected by Sir Harry 
Johnston. 

Remarks. — I have seen two other specimens, from Ankole also, 
one obtained by Sir Harry Johnston (B.M. no. 1.8.9.28) and the 
other, a young female, collected by Mr. Delme Radcliff"e (B.M. 
no. 5.4.3.4). All three are rea.dily distinguishable from the 
other grey Hyaenas of East Africa. The Abyssinian C. leo7itiewi 
is not pale grey in colour, but " braunlichgrau, auf der Mitte des 
Rlickens rostbraunlich," and has a smaller skull (zygomatic 
breadth about 155 mm., against about 165 in C. thomasi). In 
C. panganensis, from the Coast Region, the colour is da.rker, 
brownish ash, verging to rusty brown in the mane, a,nd the skull 
is considerably broader behind the postorbital processes. The 



SPOTTED HY.ENAS IN THE BRITISU MUSEUM. 99 

West- African C. togoensis appears to be the only species similar 
in colour to the Uganda B[ya?na, but according to plate 104 of 
Dr. Heck's ' Lebende Bilder aus dem Reiche der Tiere,' in 
M'hich the type of the species is figured, in the Togo Hysena the 
large spots are more regular in form, and there are among them 
some very small spots which are not present in thomasi. The 
skull of togoensis is, moreover, of the broad, not of the narrow, 
type. The original specimen is a young one, and thei-efore use- 
less for comparison, but an old topotypical specimen in the Berlin 
Museum, Prof. Matschie kindly informs me, presents the follow- 
ing dimensions : width of palate across the carnassials, 107"1 mm. ; 
upper tooth-series, 99-8; lower tooth-series, 109-3. Of course, it 
was not to be expected that a so plastic group would be re- 
presented by the same form in two countries so widely separated. 
I have much pleasure in naming the Uganda Hysena in honour 
of Mr. Thomas, as a remembrance of the kindness shown me while 
examining the invaluable collections under his charge. 

Orocuta nyas^, sp. n. 

Diagnosis. — A pale yellowish Hysena, with large dark spots and 
pale brown feet. 

Coloior. — General colour dirty bujff; mane pale ochraceous ; 
belly dark brown. The spots are dark Vandyke-brown, large and 
very scattered on the body, smaller and more close-set on the 
limbs. On the sides of the neck there are some faint traces of 
brownish-red spots. The muzzle is of a pale, dirty wood-brown, 
and the feet are of this same colour, somewhat lighter. 

Skull. — Palate long and narrow, its greatest breadth practically 
equal to the length of the upper tooth-row. Zygomatic arches 
comparatively close to the skull. 

Skull-measarements (of adult female, paratype).— Condylo-basal 
length, 260 mm, ; zygomatic breadth, 179 ; interorbital breadth, 
61; postoi-bital constriction, 48; rostral breadth on canines, 69 ; 
width of palate across the carnassials, 110 ; mandible, from, 
condyle, 190: upper tooth-series, 111; lower tooth-series, 119; 
upper carnassial, 38 x 20 : lower carnassial, 32 x 12. 

Hah. Mount Milanji, South Nyasaland. 

Type. Immature skin and skull. B.M. no. 92.8.1.5. Col- 
lected by Alexander Why tea nd presented by Sir Harry Johnston. 

Remarks. — There are in the British Museum, besides the type, 
an old female skull (92.8.1.4), measured above, and a stuffed 
specimen exhibited in the public galleries, both received from 
Sir Harry Johnston. The species, which very probably is the 
same as found in Portuguese Ea,st Africa, is readily distinguish- 
able from C. germinans, its geographically nearest ally, this latter 
being a much darker animal, wood-brown with the mane clay- 
colour. 



100 ON A VARIKTY OF PLUMAGE IX A BED GEOUSE. 

EXHIBITIONS AND NOTICES. 

December 13, 1910. 

G. A. BouLEXGER, Esq., F.K.S., Vice-President, 
ill the Chair. 

The Secretary read the following report on the additions made 
to the Society's Menugerie during the niontli of November, 
1910:— 

The nninber of registered additions to the )Society's Menagerie 
during the month of November last was 181. Of these 72 were 
acquired by presentation, 31 by purchase, 33 were I'eceived on 
deposit, 35 in exchange, and 10 were born in the Gardens. 

The number of departures during the same period, by deaths 
and removals, was 209. 

Amongst the additions special attention may be dii'ected to : — 

1 American Bison [Bison americamos), born in the Menagerie 
on Nov. 1st. 

5 Canadian 'Wi\:piii (Oerviis canadeiisls), 1 J , 4 $ , from Canada, 
purchased on Nov. 8th. . »^ 

1 Tasmanian Wolf (Thylacinus ci/nocephalas), c? , and 2 Tas- 
manian Devils {Sarcophil/cs sataiiicus). from 'J'asinania, purchased 
on Nov. 21st. 

Dr. H. Hammoxd Smith, M.Ii.C.S., E.Z.S., exhibited a mounted 
specimen of the Red Grouse {Lagopus scoticus) which displayed 
a curious variety of the ordinary plumage, and made the following- 
remarks : — 

" This Grouse was sent to me by Mr. Wynj-ard Dixon, of 
Sheffield, from the Gilkerscleugh Moors, Abington, Lanarkshii'e, 
on October 4th this year. The bird is a cock. Tlie colouring is 
not common, and the specimen is not quite like the so-called 
peppei'-aiid-salt variety, of which I saw a good example at 
Mr. Boyals, where this bird was set up. In the bird now 
exhibited, while the wings show grey colour, the feathers under 
the chin are more like the colour of the feathers of a young cock 
pheasant, a,nd there is just an appearance of a light ling round 
one side of the neck ; there are also a few bronze-coloured 
feathers on the flanks. Of this moor Mr. Dixon says in one of 
his letters : ' I was much surprised the first time I came to this 
district to find pheasants on the moors considerable distances 
from coverts or farms — in fact, the pheasants do not appear to 
come into the spinneys till November ' ; he further states that 
he has seen them two miles from any covert. But all who have 
shot on moors adjoining pheasant preserves are aware that 
pheasants will stray for very long distances over the moors from 
the coverts, especially if there a,re bilberries to be found. It has 
been suggested that this bird may be a hybrid between the 
pheasant and the grouse ; I can find ho pre\ious i-ecord of kSucIi 



THE OCCiriTAL REGTOX OF THE HEAD IX BATRACHIA URODELA. 1 01 

a. hybrid. Grouse vary considerably in their colouring from very 
dark to cream-colour. I do not personally contend that this bird 
is a hybrid ; in my opinion it is a Grouse, and I show it this 
evening as a curious variety of the colouiing of the ordinary 
Red Grouse." 

Other zoologists present confirmed Dr. Hammond Smith's 
opinion that the bird was not a hybrid. 

Mr. D. Seth-Smixh, F.Z.S., Curator of Birds, exhibited some 
slans of the Austi-alian Yellow-rumped Finch {Mnvia Jiavi- 
prymna). These birds had been kept alive in an outdoor aviary 
in England, and had developed certain markings tending towards 
those of another closely allied species, Mtmia castaneithorax. 
The exhibitor attributed this to the fact that the former species 
was a desert fox'm of the latter, and when placed in a, humid 
environment tended to revert to the plumage of the lattei-. He 
referred to a paper he had published on this subject in the 
' Avicultural Magazine,' 1907, p. 195. 



Dr. W. E. HoYLE, M.A., F.Z.S., English Member of the Inter- 
national Commission on Zoological Nomenclature, explained the 
Report presented to the Graz Meeting of the International 
Zoological Congress, and referred in particular to the proposals 
made for the protection of well knoAvn zoological names. 

A discussion followed on the portion relating .to the formation 
of an Official List of most frequently used Zoological ISTames. The 
feeling of the Meeting was very strongly in favour of the Inter- 
national Congress giving its authority to the forming of a List of 
Zoological Names, the significance of which should not be altered 
by application of the rules of the International Code. It was 
Tinanimously agreed to accept the action of the Congress if it 
would adopt this course. 



PAPERS. 

8. On the Segmentation of the Occipital Region of the Head 
in the Batrachia Urodela. By Edwin S. Goodrich. 
M.A., F.R.S., F.Z.S., Fellow of Merton College, Oxford. 

[Received November 29, 1910 : Read December 13, 1910.] 
(Text-figures 29-51.) 

rutroduGiion. 

It is now well known that in the Craniata Gnnthostomata the 
region of the head lying behind the auditory capsule is a, 
compound structure, formed of a number of segments -jriginally 



102 ME. E. S, GOODRICH ON THE OCCIPITAL REGION 

like those of the trunk. A process of cephalisation leads to the 
fusion and partial suppression of a number of skeletal segments, 
or scleromeres, which combine into a compact occipital region 
continuous with the remainder of the skuil in front. Through 
its wall issue segmental nerves. At the same time, there is a 
tendency for the corresponding muscular segments to become 
reduced. The history of this subject has been so often told that 
it need not be repeated here (SeAvertzoif 9, Gaupp 3, and 
myself 6). 

The occipital region in the Amniota has been found to include 
behind the vagus nerve four scleromeres enclosing three roots of 
the hypoglossus nerve. There are therefore probably at least 
live segments altogether between the auditory capsule and the 
atlas — the first corresponding to the glossopharyngeal, the next 
to the vagus, and the last three to the hypoglossal. Possibly 
there are a few more. 

In the Pisces the posterior limit of the head is both less 
definite and more variable in position ; but the postanditory 
region probably always includes at least seven segments. The 
Selachians have been most thoroughly studied, and in them there 
are about eight segments behind the auditory capsule (metaotic 
segments). The first corresponds to the glossopharyngeal nerve 
and the fourth mesoblastic somite (three of these being prootic) ; 
the next four segments belong to the vagus, and the last three to 
the hypoglossus, much as in Amniotes. But in the Selachian the 
anterior sclerotomes and myotomes are more distinct. The first 
metaotic somite produces no myotome, and therefore preserves no 
ventral nerve-root. The remaining occipital somites contribute to 
the hypoglossal musculature, and their corresponding ventral roots 
are the spino-occipitals of Fiirbinger (s — z). But the muscles of 
these segments tend to disappear in ontogeny from before back 
wards. In adult Selachians some two or three hypoglossal roots 
are usually found piercing the occipital region of the skull. An 
examination of the early stages, however, reveals the complete 
series of somites, nerves, and skeletal segments, which make 
up the postauditory region of the head. 

"Very different is the state of things in the Batrachia (Am- 
phibia). Here the skull appears to end immediately behind the 
vagus foramen ; and nerves contributing to the hypoglossal issue 
from the vertebral column behind the occipital condjdes. 

The first, second, and third spinal nerves may form the 
complex hypoglossal ; the second being the main, and often the 
only, hypoglossal nerve in the adult. 

These facts immediately suggest several questions : — Does the 
occipital region of the Amphibian really include fewer segments 
than that of the other Gnathostomes, or have certain segments 
been telescoped and practically crushed out ? Are the hypo- 
glossal segments of the Gnathostomes really represented by the 
first three trunk-segments of the Amphibian, or have these simply 
assumed the function originally fulfilled by others farther forward ? 



OF THE HEAD IN THE BATRACHIA URODELA. 103 

Further, if the Amphibian head includes fewer segments, it may 
be aslved whether this condition is primary, or due to the return 
of segments to the trunk which formerly held a place in the head. 

It is essential before attempting to answer these questions to 
determine how many head-segments can actually be tx-aced in 
ontogeny. Other observers have attacked the problem, but 
their results are not in agreement. With a view to settling this 
point I undertook the study of the development of the head- 
region in the Axolotl [Ambly stoma tigrinuTn). 

My method has been to reconstruct series of sections on 
paper. For this purpose it is important to have a very com- 
plete set of stages cut in various directions. I have to thank 
Dr. J. W. Jenkinson for the loan of a lai-ge number of excellent 
series of sections filling up the gaps in my own preparations. An 
appropriate number of stages has been selected for represen- 
tation in the figures given in the text ; but it will be undei-stood 
that intermediate stages have been examined. Of these figures, 
nos. 33-38, 40-43, and 45 49 were drawn from Mr. Jenkinson's 
series. 

Description. 

Before describing my own observations it will be well to give 
a brief account of the results obtained by previous workers. A 
good general description of the development of the skull of the 
Axolotl has been given by Parker (7). He describes the basilar 
plate, or floor of the cranium behind the pituitary fossa, as formed 
of a parachordal extension of the trabecular bars, combined with 
" proper occipital parachordals behind." The latter give rise to 
the occipital condyles ; but their exact relation to the myotomes 
is not elucidated, although Parker notices that the glossopharyn- 
geal ganglion lies behind the auditory capsule between the first 
and second myotomes. Stohr (11) has given a very similar account 
of the development of the skull of Siredon [Ambly stoma). Like 
Parker he found that the postpituitary region of the skull develops 
from three separate sources : the anterior parachordals (" Balken- 
platten ") derived from the trabeculae, the auditory capsules, and 
the occipital arches. By means of reconstructions of sections he 
made out clearly the origin of the posterior occipital arch. These 
arches, compared to vertebral arches, grow over the brain above, 
and along the sides of the notochord (occipital parachordals of 
Parker) join the backward extensions of the trabeculse (" Balken- 
platten "), and enveloping the notochord form the basilar plate, 
which subsequently fuses with the auditory capsules. Stohr, 
however, did not make out the exact position of the occipital 
element with regard to the nerves and myotomes. 

Sewertzoff (9) was the first author to attack this problem directly. 
According to his account there are two metaotic somites, giving 
rise to two myotomes in the embryo (text-fig. 50, B, p. 116). 
The 1st disappears later. The 2nd, corresponding to the vagus, 
remains. Behind it, in the septum between the 2nd and 
3rd, develops the occipital arch. The first trunk-segment 



104 



MR. E. S. GOODRICH OX THE OCCIPITAL REGIOX 



(3rd nietaotic) has a myotome, a ventral root, but no ganglion. 
The next and succeeding trunk-segments are complete. That 
this description is incorrect has already been suggested by Miss 
Piatt, who has given us a detailed and admiiable account of the 
development of the head in JS^ectitrus (8). 

Miss Piatt analyses the postauditory region as follows : — The 
1st somite belongs to the glossopharyngeal segment, develops no 
myotome and has no ventral root. The 2nd, 3rd, and 4th are 
vagus segments ; the 2nd somite disappears venti'ally, but its 
dorsal portion develops muscle which combines with the more 
fully formed myotome of the 3rd somite. The 4th and succeeding 
somites develop myotomes. The 3rd, 4th, and 5th grow down 
ventrally to give rise to the hypoglossal muscles, supplied by the 
ventral roots of the 4th and 5th segments. The 6th segment 
(3rd of the trunk) is the first to have a complete spinal nerve, 
with ganglion, ventral and dorsal root. The first neural aich lies 
between the 4th and 5th somites. Between the 4th and 3rd 
somites appears the occipital arch marking the hind limit of the 
skull ; while between the 3rd and 2nd somites is formed a rudi- 
mentary preeoccipital arch, which is taken into the auditory 
capsule. If this account is correct, thei-e are three metaotic 
segments in the head of Amphibia,, the myotomes of the last two 
being represented in the adult by the anterior region of the dorsal 
temporal muscle. 



mx. sop. 




hi/a. br.' ^^^^ 



Dissection of a full-grown Axolotl, showing the skeleton and nerves of the head 
and three trunk-segments. The cartilage is dotted. "View from left side. 

Coming now to my own observations on Amblystoma, we may 
begin with a glance at the structure of the full-grown animal as 
shown in text-figs. 29 and 30 *. The hyomandibular branch of the 
facial nerve issvies from behind the otic process of the quadrate 



* For explanation of the lettering of these text-figures see p. 120. 



OF THE HEAD IN THE BATRACHIA URODELA. 



105 



and the overlying squamosal, and passes down the hyoid arch. The 
glossopharyngeal and vagus come out together behind the auditory 
capsule — the former supplying the first branchial arch, and the 
latter giving off three branches to the remaining ai'ches. A large 
viscei'al branch of the vagus runs along the alimentary canal, a 

Text-fio-. 30. 




Outline of spinal cord and hind brain, with roots of cranial nerves and first tvvo 
spinal nerves, seen from left side. 

dorsal branch upwards, and slender latei'al line branches to the 
skin. Through the bony neural arch of the first vertebra issues 
the first spinal nerve, having a. ventral root only. Passing over 
the vagus it soon joins the complete second spinal, which has 
two roots *, and comes out behind the first vertebra. A dorsal 
twig of the first spinal supplies the temporal muscle. The first 
and second spinals together make up the hypoglossal nerve. 




spg 



Partial reconstruction of the hind region of the skull and the anterior region of 
the trunk of an advanced larva with a head 6 mm. long. The cai'tilaginous 
skeleton and nerves are shown through the myotomes. 

Text-figs. 31 and 32 are partial reconstructions of a larva which 
has nearly acquired the structure of the fully grown animal. 
The auditory capsule is chondrified and fused to the basilar plate, 

* Miss Piatt finds only a ventral root to this nerve in Xcctio-ifs. Thcvf cuii be no 
doubt that both roots arc present in Amb/i/sfoma as well as a ganglion. 



106 MR. E, S. GOODRICH ON THE OCCIPITAL REGION 

Text-fig. 32. . 
^r fzo. sp.'-^ nz! cpg. 




Partial reconstruction of tlie hind region of the skull and the anterior region of the 
trunk of the advanced larva shown in test-tig. 31. The mj'otomes have been 
removed. 

Text-fig. 33. 



vr. na: 




poc. 



mt 



Text -fig. 34. 




Partial reconstructions of the left side of the head region of a younger larva, 
in which a large amount of cartilage is developed. 

Text-fig. 33 shows the inner wall of the auditory capsule, and the anterior myotomes 
of the trunk through which are seen portions of the skeleton and spinal 
nerves. The vagus nerves have been cut short at their exit from the skull. 
In text-fig. 34 the skeleton and nerves are more completely shown, but the 
m\isclps have been removed. 



OF THE HEAD I\ THE BATRACHIA URODEI>A. 



107 



and the prseoccipital and occipital arches. The latter is complete 
above the brain. The second muscular segment, really the third 
myotome developed in the 4th somite, is split in two by the 
vagus. The first hypoglossal nerve comes out between the skull 
and the first neural arch. 

Text-fiii. 35. 



0£. Sp."^ 



na; zpq 




poc. gt. 



Partial reconstruction of the left side of the head region of the larva shown in 
text-fisi'. 34. Represents the same structures cut back to nearer the middle 
line, the auditory capsule and side wall of the skull being- removed and the 
vagus roots exposed. 



Text-fig. 36. 




f 



fac. 
ac 



99^' 



gv 



nch. 

Kftconstraction, seen from above, of the hind region of the skull, and anterior 
neural arches. Some roots of cranial nerves are seen on the right. 



108 



MK. E. S. GOODRICH OX THE OCCiriTAL REGION 



A larva slightly younger is shown in text-figs. 33, 34, and 35, 
At this stage the ventral portion of the 2ncl myotome is still 
preserved. It lies below the vagus between the occipital and the 
prseoccipital arches. The latter is seen to be a thickening in the 
inner wall of the auditory cajDsnle, continuous below with the 
basilar plate (text-fig. 35). A reconstruction of a rather earlier 
stage seen from above (text-fig. 36) shows the capsule beginning 

Text- fig. 37. 

spg- mt.S 




Left side view of a partial reconstruction of the anterior end. Cartilage is beginning 
to appear. The traheculfe have been cut short. The pronephric funnels are 
indicated in metaotic segments 5 and 6. The ventral portion of myotome 1 
is still present. 

Text fis. 38. 




/J 



n 



spg. na 



Text-fig. 39. 
vrf^oc. vr. 




brn. 






J 



Partial reconstructions seen from the right side. In te\-t--fig. 38 arc the m^-otomes 
and pronephric funnels. The trabecula; have been cut short. 



OF THE HKAD IX THE BATRACHIA URODKLA. 



109 



tocbondrifv behind ; tlie occipital ai'cli is fused on above the vagu.s. 
The floor of the sknll widens out considerably from the occipital 
segment forwards to allow space for the bixiin, and the preoccipital 
arch is placed opposite the point where the occipital parachordal 
plate meets the anterior pai-achordal plate, a point not yet chon- 
drified. Although it appears to chondrify in continuity with the 
auditory capsule, the prajoccipital arch is probably serially homo- 
logous with the neural ai-ches, as already suggested by Miss Piatt. 




nch 



Reconstruction, seen from above, of the licaJ and anterior trunk-region. Tlie dorsal 
ends of tlie gill-arclies are shown, also the ventral region of the myotomes. 
Cranial nerve-roots are indicated on the right side. 



Its first appearance as a procartilaginous rudiment is seen in text- 
tig. 37, taken from a still younger larva, in the septum between the 
2nd and 1st myotomes. At this stage the ventral portion of the 
first myotome (2nd somite) is still present. The auditory capsule 
itself is scarcely recognisable and quite without cartilage. 

In the next younger stage, text-figs. ,38 and 39. the pra^occipital 



110 



MR. E. S. GOOBRICH ON THE OCCIPITAL REGION 



arch has not appeared, the ventral portion of the first myotome is 
larger, a ventral nerve-root is seen supplying myotome 2, and the 
neural arches are mostly in a procartilaginoas state. 

Text-fig. 40 is a partial reconstruction of another larva of about 
the same stage, but rather younger. The ventral portions of 
myotomes 1 to 6 are shown, also the spinal ganglia. Procartilage 
vaguely indicates the position of the aiiditory capsule, the occipital 
arch is scarcely yet marked out, but the next three neural arches 

Text-jBg. 41. 




Text-fig. 42. 




Reconstructions of the anterior region, seen from tlie right side. The visceral arches 
and auditory sac are seen in text-fig. 4il ; these have been removed in text- 
fig. 42, where the nerves are shown. 

have begun to develop ca,rtilage. The first myotome is quite, 
and the second myotome nearly, cut into a dorsal and a ventral 
portion by the roots of the glossopharyngeal and vagus. The 
ventral remnant of the first myotome (2nd somite) varies con- 
siderably in development, for in the younger larva drawn in 
text-figs. 41 and 42 it is hardly distinguishable. At this stage the 
trabeculse and visceral arches are the only skeletal elements visible, 
and true cartilage can scarcely be said to have appeared. 



OF THE HEAD l.N THE BATRACHIA URODELA. 



Ill 



The next younger stage (text-fig. 43), again, shows a large 
ventral piece of the first myotome. At a still younger stage 



cfr. 



Text-fig. 43. 
f.. 3s. mtj^2 



spg. 




Left side view of a reconstruction of tlie antei'ior region. 
The visceral arches are not completed. 

(text-fig. 44), the trabeculse have not appeared and the visceral 
arches are represented by mere rods of procartilage. 



mi 



Text-fig. 44. 
gv. rnr'^^ as. f. 




mt' hrls. "^3 



Right side view of a reconstruction of the anterior end of a larva in which the 
raesoblastic skeleton is represented only by procartilaginous rod-like visceral 
arches. 

We now come to embryos without true mesoblastic skeleton. 
These earlier stages are most important in determining the number 
of postauditory segments. Text-figs. 45 and 46 are reconstructed 
from an embryo 5 mm. in length ; they both are views from 
above, but text-fig. 46 reaches farther down, so as to include 
the alinieutary canal nnd optic vesicles. Myotonies 4 and 5 are 



112 



MR. E. S. GOODRICH OX THR OCCIPITAL REGION 



clearly seen correspouding to the two pronephric funnels. 
Myotomes 2 tind 3 give off diverticula passing beliind the 4th 
branchial slit to form hypoglossal muscles. The ventral portion 
of the first myotome is some way behind the auditory vesicle and 
the rudiment of the hypoglossal ganglion. Below and in front 
of the vesicle is the hyoidean somite ; while in front of the 
rudimentary spiracular gill-slit lies the niandiljular somite. 

Text-fiff. 45. 




■ prn 



Partial vec on struct ion, seen from above, of the anterior region of an embryo in which 
the mesoblastic skeleton has not yet appeared. The epidermal thickenings 
cot responding to the glossopharyngeal and vagus nerves are indicated by 
dots. 



Another embryo of about the same length, but a little younger, 
is shown from the side in text-fig. 47. Here the auditory vesicle 
is still quite continuous Avith the epidei'mis, and the relation of the 
.somites to the gill-slits is well shown. The slits are represented 



OF THE riEAD IX THE BATIIACIIIA URODELA. 

Text-lig. 46. 



113 



mf.' 



mt. 




Similar reconstruction of the more dorsal region of the same embryo 
as in text-fig. 45. 

Text-fiff. 47. 




ocp. mds 



JJeft side view of a reconstruction of an embryo 5 mm. long. The cavities of the 

gill-pouches are shown in black. The auditory sac is a thickening directly 

\"' -continU-ouswith'the'epiblast.- Dots indicate epidei-malthickertings. .■ i-.^'- 

Proc. Zool. Soc— 1911, No. YIII. 8 



114 



MR. E. S. GOODRICH ON THE OCCIPITAL RKGIOX 



in black, though not yet open. In front is seen the mandibular 
somite, below the rudiment of the trigeminal ganglion. Behind 
it lies the hyoidean somite, just in front of the vesicle and below 
the facial rudiment. Posterior to the vesicle is the glossophar- 
yngeal rudiment, below which extends the first metaotic somite 
passing downwards into the first branchial arch. The vagus 
rudiment extends over the next two and part of the 4th somite ; 
the first myotome is placed over the 2nd branchial arch. 

Text-fig. 48 is a plan of a young embryo 3 mm. long, seen from 
the dorsal aspect, the ners^ous system being partly removed. The 

Text-fiff. 48. 




mt.3 



Partial veconstraction of the anterior region of an embryo 3 mm. long, seen from 
above. The greater part of the central nervous system has been removed to 
expose the alimentary canal, notochord, &c. The first and second metaotic 
somites overlie the first and second gill-arches. 



position of the somite with relation to the other structures is 
much the same as in text-fig. 47. Lastly, in text-fig, 49 is drawn 
the youngest stage we need investigate. The dorsal portion of 
the 2nd postauditory somite has become closely pressed on to the 
3rd somite, with which it remains intimately associated. Indeed, 
in the latter stages it is seen to be indistinguishably fused with 
it, the two combining to form the anterior region of that dorsal 
muscle which in combination with the 3rd myotome makes up 
the temporal muscle of the adult. Neither. at this nor at ajiy 



OF THE HEAD IN THE BATRACHIA URODELA. 



115 



otiier staofe does the first metaotic somite produce iimscle- 
sub.stance. 

Text-fic 49. 



mt 




Right side view of a reconstruction of an embryo 3 mm. long. The first metaotic 
somite appears below the developing anditory sac and the glossopharyngeal 
epidermal thickening. Thi 2nd somite is closely applied to the 3rd. 



Conclusions. 

From the foregoing account it will be understood that in 
almost every particular my observations confirm the conclusion 
reached by Miss Piatt in her study of Necturus. In Ambly- 
stoma as in Necturus there are three occipital segments. The 
first metaotic somite, hovi'ever, disappears very early. Probably 
the examination of an insufficiently complete series of stages 
misled Sewertzoff (9) into the belief that there are only two 
metaotic somites ; the first being either missed or later confused 
with the second. Gaupp, in his excellent review of the develop- 
ment of the skull (3), seems rather to favour Sewertzoflf's inter- 
pretation. But if the diagram given by the lattez^, and repro- 
duced by Gaupp, were correct, the second and third branchial 
rami of the vagus would lie outside the head area in the first two 
trunk-segments ; and the third bra.nchial ramus, passing behind 
the last gill-slit, would then belong to the second trunk-segment 
already provided with a complete spinal nerve possessing two 
roots and a ganglion. This is obviously not the case. The 
results of Sewertzofi" and myself are compared in text-fig. 50. 
Only in unimportant details do my own observations difier from 
Miss Piatt's. For instance, I find a ventral nerve-root in seg- 
ment 3 and a dorsal root in segment 5. Moreover, the ganglion 

8* ^ 



116 



MR. E. S. GOODRICH OX TTIK OCCIPITAL REGIOX 

Text-fio'. 50. 



poc. I 



./.v-.9-...!-,o 



■% m 



5/ m'f/ 



:(■:' ' 
I 



© 






<.y. 



Ws 



I r 



s/?^ 



B 



SjO • 



..»-:.^ -'^i 



vrt 



'■■(■iilli"i>i>i 

I 

Diagrams of the segmentation of the metaotic region in Amphibia : 
B according to Sewevtzoff, A according to my own observations. 

of seo-ment 4 seems to be early included in the rudiment of the 
vagus' ganglia. Finally, text-fig. 51 represents what I believe to 
be the true composition of the Amphibian head. 

Text-fig. 51. 




sp- hyp. 

brsl.' hypm 

Diagram of the segmentation of the anterior region of an Amphibian. 

We may now turn to the questions suggested on page 102. If 
any segments have been stippressed, it is apparently behind the 
vao-us - root that they I'lave been crushed out. Gegenbaur (4) 
befieved. the occipital arch to be « compound of several skeletal 



OF THK UEAU IX THE BATKACHIA URUDKLA. 117 

segments, not the homologue of a single neural arch a,s Stohr 
supposed. Fiirbringer adopts much the same view in his great 
monogi'aph (2), concluding that the occipital condyle of the 
Amphibia occupies the same segmental position as the last 
occipital segment in the Selachii. Many segments are supposed 
to have been crushed out between this and the vagus corre- 
sponding to the region in the Selachian skull through which 
pass the spino-occipital nerves. He claims, indeed, to have found 
possible ti'aces of these segments in C'rypioh-anchus, where he 
discovered a smalh ventral nerve-root {z.) piercing the occipital 
bone. Miss Piatt and Gaupp (8 and 3) are inclined to accept 
Fiirbringer's view. IS'ow the adoption by so many anatomists of 
the theory that a number of segments have vanished from the 
Amphibian head by a pi-ocess of excalation is, I venture to think, 
based not so much on facts as on theoretical considerations. 
In the first place, there is a reluctance to admit that a structure 
like the occipital condyle can be homologous in two groups, 
although developed in different segments. But there can be no 
doubt that the homology of an organ is independent of its 
position in the segmental series. 

The hind limbs of a Frog, an Axolotl, and an Amphiuma are 
homologous, in spite of the fact that they are placed in different 
segments. It is unnecessary here to recapitulate in full argu- 
ments which have ali'eady been given at length elsewhere (5, 6) 
with regard to the development of the fins of fishes. But I may 
briefly state : — That every trunk-segment is capable of producing 
limb-elements ; that is to saA', of contributing to the formation of 
median and paired fins. The shifting of a fin up or down the 
body is not due to the migration of fin -material from one place to 
another in the course of ontogeny ; the fin, as a whole, arises 
from that region of the trunk which it occupies in the adult (as 
is shown by the development and the nerve-supply). Relative 
displacement in ontogeny is due almost entirely to " concentra- 
tion," a relative narrowing of the base of the fin. Change of 
position in phylogeny is brought about by progressive reduction 
on one side, and growth on the other ; apparent migration is due 
to certain segments beginning to contribute to the fin at one 
end and certain other segments ceasing to contribute at the 
other. By such " transposition " the fin may reach an entirely 
new position. 

Kow, in the case of the fins of fishes, I have already shown (5) 
that it is not possible to account for variation in position by 
the theoiy of inter- and excalation. Growth and transposition 
from one segment to another alone account for the facts. The 
same is probably true of the occipital condyle. Thei'e is not the 
slightest trace of the disappearance of segments behind the vagus 
in the ontogeny of the Amphibia. We are familiar with tiie 
variation in the extent of the gill-region in Yertebratesby mei'e 
growth. Obviously tlie hind limit of the series of gill-slits varies 
1)ack\\"ard . or forward, according •. as cLTtain.: segmcuts. cease'.tu 



118 MR. E, S. GOODRICH OX THE OCCIPITAL REGION 

develop gills or take on the function of gill -formation. The 
posterior limit of the skull is doubtless altered in the same way, 
and the position of the occipital condyles may shift vip or down 
the segmental series. There should, therefore, be no theoretical 
objection to accepting the ana,tomical and embryological evidence 
that the occipital region of the head in Amphibia contains only 
three segments. If segments conld really disappear, leaving no 
trace behind, it would be hopeless to attempt to homologise 
segments in any two forms. 

There is another theoretical consideration which seems to have 
led to the adoption of the view that the occipital region of the 
Amphibia is not as simple as it appears. It is urged that if it 
contained only three segments, the Amphibia would be more 
primitive than the Fishes from which they have descended. The 
possibility of the reduction in the number of occipital segments 
has just been explained above ; but is there really any necessity 
to assume that it has taken place ? From whatever Fishes the 
Amphibia may have been derived, we may be sure it was not 
from fvilly specialised Selachians. Fi-om palaeontology alone we 
may hope to obtain definite evidence on this point ; until con- 
trary evidence is brought forward, there is no necessity to assume 
that the ancestors of the Amphibia had more than three differen- 
tiated occipital segments. Of all the living fishes the Dipnoi are 
those which most closely approach the Amphibia ; even in the 
modern Ceratodus there is no occipital condyle, no distinct limit 
between head and trunk. It is true that several trunk-segments 
have here been more or less completely assimilated to the skull 
(Sewertzofif, 10) ; bvit thei-e is no reason to believe that in the 
remote common ancestor of the Dipnoi and Amphibia the dividing- 
limit between the two regions could not have been developed 
three segments behind the auditory capsule. This seems to be 
in agreement with the conclusions of Agar (1), based on a study of 
the development of Lepidosiren and Protopterus '*. 

The hypoglossus in the Amphibia and Amniota may certainly 
be considered as homologous, although not necessarily composed 
of the same segmental nerves. It is owing to the shortness of 
the skull in the Amphibian that the hypoglossal roots come out 
behind it. 

Summary. 

The chief contents of this paper may be summarised as follows. 
Three occipital segments occur in tlie head of Amhlystomu. Of 
the three somites developed in these, the fii-st forms no muscle 
and disappears early ; the second forms a myotome divided into 
dorsal and ventral portions, of which the former alone persists, 
fusing with the myotome next behind. The myotome of the 
third segment persists dorsally, that of the first trunk-segment 

* The occipital region in fossil Amphibia seems to have been formed as in the 
modern species. Important evidence as to the assimilation of the hypoglossal 
segments in primitive Amniotes may perhaps be gleaned from a careful investiga- 
tion of early fossil Reptiles. For instance, it seems to me not improbable that in 
FareiasaHVus the occipital region is still in nn intermediate condition. 



OF THE HEAD IN TUE BATRACHIA URODELA. 119 

combines with it to form the temporal longitudinal muscle 
supplied in the adult by the first spinal nerve. The glosso- 
pharyngeal nerve belongs to the first metaotic segment and has 
no ventral root. To the second segment belongs the vagus root, 
with which seems to be combined the ganglia of the next two 
nerves. No ventral root was found in the second segment ; but 
a ventral root occurs in the last metaotic segment in early stages 
of development. The first spinal issues from between the skull 
and first neural arch as a ventral root only ; it joins the second 
spinal to form the hypoglossal nerve supplying muscles derived 
from ventral outgrowths of the second, third, and fourth myotomes. 
The basilar plate of the skull is formed by a backward growth of 
the trabeculae meeting a forward growth of the base of the 
occipital arches. It fuses with the auditory capsules, and with 
the prseoccipital arches developed in the septum between the first 
and second myotomes. The occipital arches arise in the septum 
between the second and third myotomes. There is no evidence of 
the disappearance of segments behind the vagus, and no valid 
objection to the view that the hind limit of the skull may shift 
backwards or forwards in the course of phylogeny. At the same 
time there is no reason to suppose that the ancestors of the 
Amphibia had more than three occipital segments, when the 
occipital condyle became clearly defined. 

List of References. 

1. Agar, W. E. — " Development of the Anterior Mesoderm 

and Paired Fins in Lepidosiren and Protopterus." Trans. 
Roy. Soc. Edinburgh, vol. xlv. 1907. 

2. FuRBRiNGER, M. — " Ueber d. spino-occipitalen Nerven." 

Festschr. v. 1. Gegenbaur, vol. iii. Leipzig, 1897. 

3. Gaupp, E. — " Die Entwicklung des Kopfskelettes." Hert- 

wig's Handb. Entw. Wirbeltiere, vol. iii. Jena, 1906. 

4. Gegenbaur, C. — " Die Metamerie des Kopfes." Morph. 

Jahrb. vol. xiii. 1888. 

5. Goodrich, E. S. — " On the Development &c. of the Fins of 

Fish." Quart. Journ. Micr. Sci., v. 1906. 

6. Goodrich, E. S. — The Vertebrata Craniata : Cyclostomes 

and Fishes. Treatise on Zoology : part 9. London, 1909. 

7. Parker, W. K. — " On the Structure and Development of 

the Skull in the Urodelous Amphibia." — Pai-t I. Phil. 
Trans, vol. 167. 1876. 

8. Platt, J. B. — " The Development of the Cartilaginous Skull 

and of the Branchial and Hypoglossal Musculature in 
JVecturus" p. 377. Morph. Jahrb. vol. xxv. 1896-8. 

9. Sewertzoff, a. N. " Die Entwicklung des Selachierschadels."' 

Festschr. 0. v. Kupfi'er. Jena, 1899. 

10. Sewertzoff, A. N. " Zur Entwicklung des Ceratodus forsteri'' 

Anat. Anz. vol. xxi. 1902. 

11. Stohr, Ph. — " Zur Entwicklung des Urodelenschadels," p. 477. 

Z. w. Zool. vol. xxxiii. 1880; "Zur Entwicklung des 
Anurenschiidels." t), 68. [hid. vol. xxxvi. 1882. 



120 



MR. OLDPIELD THOMAS OX THE MAMMALS- 



Explanation af Lettering on I'ext-figures. 



a. 


aiiditory nerve. 


ma., 


ac. 


auditory capsule. 


md., 


acp. 


ascending process of quadrate. 


mds., 


as. 


auditory sac. 


Vlt., 


hp., 


branchial plexus. 


mtJi., 


br. 


branchial branch of vagus. 


mx., 


hrii. 


brain. 


na. 1, 


brs. 


mesoblast of branchial arches 


ncJi., 




1-4. 


ns.. 


brsl. 


branchial slit 1-4- 


oc, 


emm. 


median mandibular cutaneous 


OCp., 




branch of facial. 


opn.. 


col. 


columella auris. 


per., 


ctr. 


crista trabeculie. 


poc, 


e. 


eye. 


pop.. 


et. 


epiblastic thickening. 


pvf.. 


/, 


facial nerve. 


prn.. 


fac, 


floor of auditory capsule. 


2-> 


foe, 


oculomotor foramen. 


s. \, 2, . 


foP' 


optic foramen. 


Sll., 


9f; 


facial ganglion and its epiblastic 


sJc, 




thickening. 


sm.. 


ggi; 


ganglion of glossopharyngeal. 


sop.. 


9I; 


glossopharyngeal nerve and its 


sp.l. 




epiblastic thickening. 


sp.i!d& 


gt., 


glottis. 




gtr., 


ganglion of trigeminal and its 


spg-, 




epiblastic thickening. 


spl., 


g"; 


ganglion of vagus and its epi- 


spvr.. 




blastic thickening. 


tr.. 


h.. 


hyomandibular branch of facial. 


trg.. 


Jida., 


hyoid arch. 


trp., 


hyp., 


hypoglossal nerve. 


tso., 


Tiypvi., 


hypoglossal muscle. 


■f'-j 


im.. 


internal mandibular branch of 


vll.. 




trigeminal. 


V7-., 


J-> 


anastomosis of Jacobson. 




11., 


lateral line nerve. 


■vrf.. 


■m.. 


medulla. 





mandibular arch. 

mandibular branch of trigeminal. 

mesoblast of mandibular arch. 

myotome 1-6. 

mouth. 

maxillary branch of trigeminal. 

neural arch. 

notochord. 

nasal sac. 

occipital arch. 

optic cup. 

optic nerve. 

pericardium. 

preoccipital arch. 

preorbital process. 

profundus nerve. 

pronephros. 

quadrate. 

metaotic somite, 
hyomandibular somite, 
hind limit of skull, 
mandibular somite, 
superior ophthalmic nerve, 
first spinal lierve. 
f, second spinal nerve, dorsal and 
ventral root, 
spinal ganglion, 
spiracular slit, 
ventral i-oot of spinal nerve, 
trabecula ci'anii. 
trigeminal ganglion, 
trabecular parachordal, 
tectum synoticum. 
vagus nerve. 

ventral lateral line nerve, 
roots of vagus and glossopharyn- 
geal, 
ventral root. 



9. The Mammals o£ the Tenth Edition o£ Liunsens ; an 
Attempt to fix the Types of the Genera and the exact 
Bases and Localities of the Species. By Oldfield 
Thomas, F.R.S., F.Z.S. 

[Received November 29, 1910: Eead December 13, 1910.] 

The tenth edition of Linne's 'Systema Naturae' is the recognized 
basis and commencement of all systematic zoological work, but 
doubtful questions in its interpretation are very numerous, and 
there is probably nothing more desired, by mammalogists at least, 
than a methodical examination of this important work, with 
suggestions, obtained on some definite and uniform system, foi- the 
identification of its types, both of genera and species. 

For many years I have taken a very great interest in this 
subject, a,nd have now ventured to prepaie the present paper, 
with the hope that it may be of service to other mammalogists less 
favourably .situated as regards book.s and specimens than 1 have 



OF THE TKNTH EDITION OF LINX.EUS, 121 

the privilege of being. For iu all the work I have scarcely in 
a single instance been vinable to obtain from the library of the. 
Natural History Mvisenm what books I have wanted to verify 
Linneeus's quotations, however rai-e and obscui'e they have been. 
Without such an advantage it would have been impossible to 
do the work at all. 

With regard to the method used for finding the type-species of 
the Linnean genera, I would di'aw attention to the very consider- 
able extent to which the principle of tautonj^my is available. In 
ai. great propoi'tion of cases Linnseus selected as a generic name a 
term which had been used by eai-lier writers for the single name 
of a member of the genus, so that among the species of any given 
genus there is commonly one which has among its synonyms the 
name used by Linnaeus for the genus. 

These names are generally quoted from Gesner, and comprise 
the most conspicuous and best-known members of each genus, so 
that in nearly all cases the genotype selected on this system is the 
very one which by general consent we should wish to have for the 
type. Any other rule that has been suggested, either "first 
species " or " elimination," is liable to produce results by no means 
so satisfactory — indeed, the many objections to both are too well 
known to need any insistence upon here. Nor can any later and 
arbitrary selection of types without reason assigned be taken as 
necessarily valid. 

In two cases only, those of Simia and Dasypus, are results 
arrived at difierent from those obtained by previous wi'iters. The 
shifting of the latter name from the rarer '•'■ Eujihractihs'^ to the 
really common Tatusia is unfortunate, but is necessary if we are 
to get the benefit of the general stability which is obtainable from 
the acceptance of the principle of tautonymy. 

The case of /Simia, however, is very different, and I confess to. 
hoping that the suggestions which are now being made for some 
overruling decision in the case of certain well-known but 
technically misapplied names may be carried out, both a,s to a 
number of such names in general and to Simia in particular. 

But it seems advisable, nevertheless, to put the technical case 
irrespective of the prospect of any such decision, and therefore 
(while hoping that a fiat attaching Simia to the Orang may go 
forth) I have to point out that this name should be applied to 
the Macaques, with S. sylvana (the Barbary Ape) as its type, 
while Anthropopithecus will remain available for the Chimpanzees. 

Should no such fiat as I speak of ever come to pass, this allo- 
cation of Simia is at least better than its application to the 
Chimpanzees. Not only is it less confusing in itself, owing to the 
consequent total abolition of the combination Simia satyribs, con- 
nected for a century with the Orang, but now sought to be applied 
to the Chimpanzee, biit also the genus to which it goes, that of the 
Macaques, is already ma,rked as the victim of a. noraenclatural 
change, for Pithecns antedates Macaca, and therefoi'e one change 
is not much worse than the other. . . - .... 



122 MR. OLDFIELD THOMAS ON THE MAMMALS 

Moreover, the Macaques are far less often dealt with or spoken 
of by outside writers, not up to date in nomenclatural techni- 
calities, than are the Anthropoid Apes, among whom such a change 
has the maximum of inconvenience. 

[After the completion of the present paper, there has come into 
my hands a copy of the " Opinions i"endered by the International 
Commission on Zoological Nomenclature," dated July 1910, but 
received by members of the Gratz Congress only at the end of 
October. 

In this publication the question of Linnean Genera in general 
Hnd Simla in pai-ticular is dealt with, certain conclusions are 
come to not differing widely from my own, and the principle of 
tautonymy as applied to the names in Linnteus is on the whole 
approved of. 

Two genera {Mas and Camelus) have types definitely fixed for 
them, twenty are mentioned as " seeming to retain as types " the 
species I fix for them by tautonymy, though this is " not a ruling 
by the Commission," and Dasypus is ignored. 

With regard to Simla, the Commissioners, while seeing that 
the rule brought the type on to sylvanus, would appear to have 
thought it a benefit for zoology that satyrus should be considered 
as such, an opinion in which I believe but few zoologists on 
careful consideration would be found to agree with them. Under 
this idea a brief but rather far-fetched and unconvincing argument 
is adduced to avoid the simple conclusion about sylvanus, viz. : — 
that " an examination of Gesner's text shows that he did not 
use Simla in the specific sense of ' the simia,^ and that therefore 
Siinia Gesner, as used in the synonymy of S. sylvamcs, cannot be 
considered as the name of a species." 

ISTow I have carefully examined Gesner (three editions, including 
that qvioted), and can only say that his use of Simla is exactly 
the same as his use of other names which are accepted as tau- 
tonyms. On p. 847 he writes " de Simia," and gives a figure of 
the Barbary Ape, and then on p. 855 he begins on other monkej's 
under the heading " de Simiis diversis." Exactly in the same 
way we have " de Mure" on p. 714 {Mus muscuhts) and " de 
Muribus diversis" on p. 731; " fZe Mtistela" on p. 752, '' de 
Mustells diversis " on p. 762. 

It is true that the Commissioners do not definitely accept as a 
ruling of the Commission the above-mentioned twenty names on 
the rigid basis of tautonymy, but seem to take as valid Palmer's 
selection of the types as published in his Index (1904). 

But the latter work cannot be accepted as that of a " reviser " 
in the true sense of the term. It is merely a bibliographical work 
giving an indication of what species have been currently accepted 
as types, without special revision by authors with all the facts 
before them. In no case could I admit that it should oveixide the 
clear case for tautonymy presented by Simia equally with the 
twenty genera in which its results are agi'eed to both by Palmer 
and the Commissioners ] 



OF THE TEXTH EDITION OF LTNX.*:US, 123 

The question of the species is not so simple as that of the 
genera, having been complicated by the pages and pages which 
commentators have thought it necessary to write balancing the 
pros and cons in each case, weighing the importance of this 
reference or that, and deciding that such and such is the " primary 
quotation " and should be taken as the basis of the Linnean 
name. This balancing method, however charming an exei'cise of 
the writer's ingenuity it may provide, is unfortunately full of the 
pei'sonal equation, and therefore objectionable, while any method 
which Avoi'ks automatically should be welcomed by those desirous 
of coming to a final and impersonal decision on such points. 

The system I advocate is that Linnseiis's quotation of his own 
earlier writings should be given an absolutely overriding im- 
portance, and that, where these exist and can be traced back, all 
others should be ignored. 

This principle is not an arbitrary one, but is quite reasonable. 
For when in 1758 Linnseus gave a binomial name to an animal to 
which he had already given in 1748 a species-number and a. 
diagnosis (quoting it in the tenth edition), his idea of the species 
would clearly have been formed at the earlier date, the later 
edition only adding the binomial name. To accept as of primary 
importance in the determination of the latter some reference 
which was not in existence at the time of the earlier edition 
appears to me little short of ridiculous. 

But if we take Linnaeus's own Linnean quotations — which are, 
in most cases, of the sixth edition — we get our field of selection 
narrowed down at once to one or two easily traceable references, 
from which the basis of the author's original idea of each species 
is I'eadily obtainable. 

So important did Linn?eus himself think the sixth edition that 
practically every species of Mammal in it is quoted in the tenth, 
and that merely by its page and number, as if that were the one 
edition to which attention should be paid. 

Curiously enough, this does not seem to be the case in other 
groups than Mammals, the references to the sixth edition being 
omitted, and primary stress laid on the ' Fauna, Suecica.' 

With regard to type localities, I have made every effort to 
identify the places from which the specimens came that were the 
original bases of Linn?eus's names. His own statements of 
habitat ai'e for the most part too vague to be of any use, while 
accepting them literally would sometimes lead to grotesque errors, 
such as the assertion, based on Linnseus's ''Hub. in Indiis" that 
" some Indian Armadillo " (!) should be taken as the basis of 
Dasypus septemcinctus. " In Indiis," in fact, hardly amounts to 
more than a statement that the species is exotic. 

But many localities can be settled from the authors quoted, as, 
for example, in the case of Marcgrave, on whose descriptions * 
Linnjevis's names for a large number of the commoner Brazilian 

* ■ HistoriiT Rcrum Xaturalium/ Braft.ilitc, 16i8. 



124 MR. OLDFIKLU THOMAS ON THE MAMMALS 

species were based. With all these we are able to fix as a type 
locality Pernambuco, for Marcgrave stayed at " Moritzstadt," now 
Recife, most of his time, 1640-1644, while even his excursions 
were limited to the coast region between 5° 45' and 1 1° 1 1' S., thus 
taking in little more than from Rio Giunde do Norte to Alagoas, 
a region at the centre of which Pei-nambuco lies. 

Agaiii, with regard to species named fi'om the iigures in Seba's 
'Thesaurus,' it would not be unjustifia,ble to suggest tha,t in the 
case of all tropical South American animals, Surinam — the great 
source of all Dutch collections—should be accepted as the type 
locality, except when the presei'vation of the type specimen or 
some incompatibility in the description shows this to have been 
impossible. 

The alterations in nomenclature that have proved to be necessary 
are fortunately comparatively few, certainly far fewer than 1 
ha,d originally feared would be the case. Those not previously 
published are (he following :^ — 

Simaa. Genotype not sati/rus but sylvanus. p. 125. 
Anthropopithectcs satyrtis for Chimpanzee, p. 125. 
Technical name of Barbary Ape, Simia sylvana. p. 125. 
Fygathrix a distinct genus from Preshytis. p. 127 (footnote). 
Macaca syrichta for M. philippinensis auct. p. 129. 
Mania pentadactyla for M. dahnani iiund. p. 133. 
Dasypus for Tatu; Eiiplwactus for '■'■ Dasypus" p. 141. 
Arvicanthis striatus for A. jndchellus Gray. p. 148. 
Pftyseter catodon for Ph. macrocephcdus Linn. p. 157. 

Other names here confirmed, which a,re unfamiliar, though 
not now published for the first time, are those indicated for the 
Whalebone Whales by Mr. F. W. True, those of certain monkeys 
(Oebus, Mandrill, &c.), recently pointed out by Dr. Elliot, Preshytis 
aygula for P. m.itratios, Cynocephalus t^olans for the Philippine 
Colugo, Elej)has maximus for the Indian Elephant, and Myrmeco- 
phaya tridactyla for M. juhata. 

In one respect the pi-esent paper does not complete the exami- 
nation of all the Linnean 1758 species, for a certain number are 
still unidentified pending the study of typical specimens formerly 
in the " Museum Regis Adolphi Friderici,"and no doubt in many 
cases still in existence. I would commend to my colleagues iri 
Sweden an examination of the old collections there preserved, 
with an identification of such types as are still extant. Perhaps 
Dr. Einar Lonnberg may be able to forward this final completion 
of the study of the " Zoological Bible " written by his immortal 
compatriot. 

In the following pages the genera and species are ari-anged 
as in the ' Systema,' while opposite the name of each species on 
the right-hand side of the page I have placed" the name of the 
recognized modein genus into which that species falls. 



OF TflK TEXTH KUITIOX OF LIXX.F.US. 125 

Genus 1. Homo. p. 20. 
Two species. Type, //. sapieus. 

Species : 

1. sAPiEX.s. p. 20. Homo. 
" JSTosce te ipsum." 

2. troglodytes, p. 24. 

A mixture of the abnormal hairy woman iigured by Bontius 
(Hist. Nat. Med. Ind. Or. p. 84) witli an account of albino negroes 
taken from Nils Matson. 

Genus 2. Simia. p. 25. 

Twenby-one species. Type by tautonymy : 8. sylvanus 
(" Simia," Gesn.). 

The species are divided into three sections, (1)* Cauda nulla : 
Simife veterum, (2) *"* Cauda abbreviata : Papiones, and (3) *** 
Cauda elongata : Cercopitheci. 

From the name " Simife," the type must clearly be sought 
among the first section, which contains two species, satyrus and 
si/lvamis, and the second of these [not the first) should be taken 
as type, because among its synonyms — the first, in fact — occurs 
^' Simia, Gesner, p. 847" (of 1620 edition, 957 of 1551 edition, p. 1 
of German edition). 

From this it would follow that, failing a fiat on the subject, 
the generic name Simia should be ti'ansferred — not to the Chim- 
panzee, as has been done by Matschie aiid Rothschild, but to the 
Bai-bary Ape and its congeners. 

Species : 

1. SATYRUS. Anthi'opo2nthecu3. 

Linnean reference : Syst. Nat. (6) p. 3, no. 1, where the first 

quotation is 
. Tulp. Obs. p. 270, pi. xiv. 
Clearly the Common Chimpanzee*, whose name (again failing a 
fiat) would thei-efore ))e Anthrojiopithecus satyrics. 

2. SYLVANUS. p. 25. (Simia or) Macaca f. 

Linnean reference : Syst. Nat. (6) p. 3, no. 3, where the sole 

reference is to 
Alpin. Aegypt. p. 241, pi. xvi. 
This is clearly a Barbary Ape, to which therefore, instead 

* Of. W. Eothschild, P. Z. S. 1904, ii. p. 420. 
- t I continue to use Macaca provisionally, pending a decision as to " fiat "names. 
To use Pithecus (although it antedates Macaca) would be merely the commence- 
ment of a useless familiarity with that name, which would be again overthrown 
should the tisit'principle fail. 



] 26 MR. OLDFIELD THOMAS ON THE MAMMALS 

of Macacus (or Macaco) hi.uus or Initus ecaudatus, the name 
Simia sylvana would appear to be technically applicable. 

The reference Simla sylvanus has been already frequently put 
among the synonyms of Afacaca imia, so that the present identi- 
fication is not a new one. 

3. SPHINX, p. 25. (Mandril *.) 

Linnean reference : Syst. Nat. (6) p. 3, no. 5, where the sole 

reference is to Ray (Quadr. p. 158), who in turn speaks 

of the " Baboon Gesnero descripta et depicta." 

This is, no doubt, as shown by Dr. Elliot t, the Mandrill, of 

which a figure and descrij)tion are given on the page between 157 

and 158 of Gesner's German edition (1606). 

4. APEDIA, p. 25. ('0 
Linnean (and sole) reference : Amoen. Acad.i. p. 278 (1749). 

Not determinable from descrij)tion, but if the specimen de- 
scribed in the ' Amoenitates ' as being in the Museum Adolpho- 
Fridericianum still exists, the species may yet be identified. 

Type locality " in Indiis," which means hardly more than 
" foreign," No locality at all is mentioned in the ' Amoenitates.' 

5. siLENUs. p. 26. Macaca. 

No definitive reference, the only one, that to Alpinus (p. 242), 
being with a query, so tha,t the account should be treated by its 
own evidence only. 

The description and locality, however, appear to me to be 
sufficient to assign the name to the Malabar " Lion-tailed Monkey 
of Pennant," as was done by pi'actically every author up to 
Blanford. 

That author, working on the basis of the 12th edition, first 
stated + that the name did not refer to the Lion-tailed Monkey, 
and then later on used it for it §. 

No exa.ct typical locality available. 

6. FACJNUS. p. 26. (?) 

Linnean reference: Syst. Nat, (6) p. 3, no. 15, whence we 

get Cercopithecios barbiztros, no. 1, Clusius, Exot. p, 371 

(1605), 

It does not seem possible to determine this monkey with any 

certainty. In the pointed white beard and white chest we have 

something which suggiests the Diana Monkey, while, on the other 

hand, by the length of the beard (if mantle- hairs were mistaken 

for a beard) and the tufted tail a Colobus of some sort might be 

indicated. 

No locality given. 

* As Palmer has shown, this is the technical name for the Mandrill, if considered 
generically distinct from other Baboons, as it apparently should be. 
t Ann. Mag. N. H. (8) iv. p. 417, 1909. 
I P.Z.S. 1887, p. 620. § Mamm. Ind. p. 16, 1888. 



OF THE TENTH EDITIOX OF LIXX/l':US. 127 

7. PANiscus. p. 26. Ateles. 

Linnean reference: Syst. Nat. (6) p. 3, no. 14, whence 
Guariba, Marcgr. Bras. p. 226. 
A teles paniscus. 
Type loca,lity : Pernanibuco. 

8. DIANA, p. 26. Cercojyithecus. 
Linnean reference: Act. Stockholm, 1754, p. 210, pi. vi. 

Here a full description of the monkey is given, and, as Jentink *' 
has pointed out, it is the red-thighed form to which the name 
diana was originally applied. 

Type locality : " Guinea." — Liberia (Jentink). 

9. CEPHUS. p. 27. Cercoplthec'us. 

jSTo Linnean reference. Quotations of Marcgrave (p. 228) 
and Ray (p. 156). 
lYpe locality : " Guinea." 

10. AYGULA. p. 27. Preshj/itsf. 

No Linnean reference. Sole basis : "Osb. iter. 99 "( = Osbeck, 
Ostindisk Resa, p. 99, 1757), 
Type locality : Java. 

The species usually known as P. mitratus Eschsch. (see Thos. 
k Wrought. P.Z.S. 1909, p. 373). 

11. HAMADRYAS. p. 27. Pajiio. 

Linnean reference : Syst. Nat. (6) p. 3, no. 7, where the sole 
reference is to Alp. Aegypt. p. 242, pis. xvii. & xviii, 
Hamadryas Baboon. 
Type locality : Egypt. 

12. JACCHUS. p. 27. CalUthrix. 
Linnean reference: Syst. Nat. (6) p. 3, no. 12, whence Catrui 

minor, Marcgr. Bras. p. 227 (misquoted 228). 
Common Marmoset. 
Type locality : Pernambuco. 

13. (EDIPUS. p. 28. Leonlopithecus. 

No Linnean reference. Sole basis : Edwards, Aves, iv p. 195 
pi. 195, 1751. 

Type specimen said to have been brought from " Yera Cruz, in 
New Spain," but the species does not occur there. 

* Notes Leyd. Mus. xx. p. 233, 1898. 

t Dr. Elliot (Bull. Am. Mus. N. H. xxviii. p. 151, 1910) has recently pointed out 
that the name Ft/gathrix, hased on Simia nemcea, antedates Freshytis, and should 
be used instead of it. But an examination of the skulls convinces me that nemcea 
and its close ally nigripes are, as Gray said, generically distinct from all the other 
Langurs, which would therefore retain the name Freshytis. The basal axis of the 
braiu-case is set on to the facial bones at quite a different anffiein the two genera, 



128 MR. OLDFIELD THOMAS OX THE MAMMALS 

14. ^THiops. p. 28. Cercojntkecus. 

Linnean reference : Linn, in Hasselquist, Iter, p. 190, 1757. 
Formei'ly known as C. griseovw-idis. 

See de Winton, in Anderson, Mamm. Egypt, p. 15, 1902. 
Type locality : Upper Egypt. 

• 15. MIDAS, p. 28. Leontopithecus. 

No Linnean quotation. First and primary reference : 
Edwards, Aves, iv. p. 196, pi. 196, 1751. 
Locality: "West Indies." 

16. CYNAMOLGOS *. p. 28. Pcipio. 

Linnean references: Syst. Nat. (6) p. 3, no. 10 (ex Marcgr. 

Bras. p. 227), and Simia aegyptiaca, Linn, iti Hasselq. Iter, 

p. 189, 1757. 

Both these references being Linnean, the one that refers to an 

actual specimen should be taken in preference to that merely 

based on Marcgrave's short and indefinite notice. 

Consequently Simia cyyiamolgos becomes a synonym of Papio 
hamadryas. 

Type locality : Upper Egypt. 

17. APELLA. p. 28. Cehus. 
Linnean reference: Mus. Ad. Frid. p. 1, pi. i., 1754. 

Cehus capucinus auctoi-um. 

■ Bee Elliot, Bull. Am. Mus. N. H. xxvi. p. 227, 1909. 

No type locality available, but Guiana may be accepted from 
later authors. 

18. MORTA. p. 29. Saimiri. 
Sole basis : Seba, Thes. i. p. 52, pi. xxxiii. fig. 1, 1734. 

Clearly a Squirrel-Monkey, but as the species is not determinable 
the name must lapse. 

The suggestion that Sui'inam should be taken as a typical 
locality for Seba's animals cannot be adopted in this case, as the 
statement that Simia morta had the back of the head blackish is 
inconsistent with the condition found in the Guianan species. 

■ 19. CAPUCiNA. p. 29. Cehus. 

Linnean reference: Mus. Ad. Frid. p. 2, pi. ii., 1754. 

The monke}' usually known as C. hijpoleucv,s Humb. 
: See Elliot, Bull. Am. Mus. N. H. x'xvi. p. 227, 1909. 

No type locality available. The species is Central American 
and Colombian. 



being far more strongly inclined in Fy<)athrix, in which, as a consequence, the 
posterior nares are of enormously greater height. Gray, following Dahlbohm, 
separated the two on external characters and the proportions of the fore and hind 
limbs, and my comparison of the skulls quite confirms bis judgment in the matter. 

■■*' Spelling' altered fo (;7/«())»o7^9«-s in 12tb edition.' ' ' ' ' ■'■ '" ''■ ■• '■■^■Mi''i 



OK THE TEXTH EDITION OF LIXN'.EUS. 129 

20. SCIUREA. p. 29. Sainniri. 
Linnean reference : Mns. Ad. Frid. i. p. 3, 1754. 

Common Squii^rel-Monkey. 

The Guianan form, which agrees with the description, may be 
accepted as typical. 

21. SYRICHTA. p. 29. Macaca. 

Linnean reference : Syst. iSTat. (6) p. 3, no. 6 ; whence we get 

" Cercopithecus Luzon, minimus D. Camel," Petiver, Gazyo- 

phylacium, pi. xiii. fig. 11. No. 176 of Index. 

This figure — although a ghastly travesty in itself — can be 

nothing bat the one monkey found in Luzon, viz., Macaca 

philippensis, philippinensis, or palpebrosus auctorum, which names 

should give way to that of M. syrichta. 

Type locality : Luzon (probably Southern Luzon, whence 
Dr. Camel sent " Lemur volans"). 

Genus 3. Lemur, p. 29. 

Three species, no one with " Lemur" as a subsidiary name. 

Type, L. catta ; L. tardigradtis having been made the type of 
Loris, Geoff., 1796, and L. volans of Cynocejjhalus, Bodd., 1768 
(= Galeopithecus^ Pall., 1780). 

All authors have accepted L. catta as the type of Lemur. 

Species : 

1. TARDIGRADUS. p. 29, Loris. 

Linnean references : " Mus, Ad, Fr, i, p, 3 ; Syst. Nat. (6) 
p. 5 *, no. 2." 

The specimen referred to in Mus. Ad. Frid. and redescribed in 
the 10th edition is still in the Stockholm Museum, and, as 
Dr. Lonnberg informs me, is the Cinghalese Slender Loris. It 
may even have been the specimen figured by Seba, who is quoted 
in the 6th edition. 

See also Thos. Ann. Mag. N. H. (8) i. p. 467, 1908, where the 
same conclusion is reached on the ordinary " Linnean reference " 
grounds. 

Type locality : Ceylon, 

2. CArrA. p. 30. Lemur. 

Sole basis : Simia — Sciurus madagascariensis, sive Maucauco, 
Edwards, Aves, 119, t. 199. 
The Ring-tailed Lemur. 

H. VOLANS. p. 30- Cynocephalus. 

References : — 

Cato — Simius volans camelli, Pet.(iver), Caz.(ophylacii 
Naturfe et Artis), t. 9. f. 8 (1702); Act. Angl. 277, 

* Misprint for .3, 

PB.oa Zooh. Soc- 1911, No. IX. 9 



130 Ml!. OLUFIELD TliOMAS ON THE MAMMALS 

p. 1065 (= Phil. Trans. Roy. Soc. pt. 277, included in 
vol. xxiii. 1704). 
Yespertilio admirabilis, Bont.(ius), Java, p. 68, 1658. 
Felis volans ternatia, Seba, Mus. i. p. 93, t. 58. f. 2, 3. 
The first is the primary and only vital reference, as the others 
refer to specimens from impossible localities (Guzerat and Ternate). 
Petiver's type locality was Pampanga, Southern Luzon, and the 
animal therefoi^e was that until i-ecently known as G aleopithectis 
(or Colugo) philippinensis. 

Cynocephalus, Boddaert, was based on " Lemur volans Linn.," 
and antedates Gcdeopithecus hj many years. 

See Thos- Ann. Mag. N. H. (8) i. p. 252, 1908. 

Genus 4. Vespertilio. p. 31. 

Seven species, of which no. 7, murimos, has among its synonyms 
Vespertilio, Bell. Av. 147; Gesn. Av. 694. 

Therefore type by tautonymy : Vespertilio mnrinus L., the bat 
formerly known as F. discolor. 

See Miller, Ann. Mag. N. H. (6) xx. p. 379, 1897. 

Species : 

1.. VAMPYRUS. p. 31. Fteropus. 

Linnean reference : Syst. Nat. (6) p. 7, wo. 1. 

There the only reference is to 
Canis volans ternatanus orientalis, Seba., Thes. i. p. 91,- 1. 57. 
f. 1, 2. 
Fteropxis edidis auctorum. 
See Thos. P. Z. S. 1892, p. 316. 
Type locality : Ternate. A co-type in British Museum. 

2. SPECTRUM, p. 31. Vampyrus. 

Onlv reference : Canis volans maxima auritii, Seba, Thes. i. 
p". 92, t. 58. f, 1. 
Species known rightly as Vampyrus spectrum L. 
Suggested type locality : Surinam. 

3. PBRSPiciLLATDS. p. 31. Hemid^erma. 
Linnean references : Syst. Nat. (6) p. 7 ; Mus. Ad. Fr. i. 

p. 7. 

The latter, as referring to an actual specimen, would take 
precedence, but the description is unrecognizable, and no specimen 
is now in existence, as Dr. Lonnberg informs me. We therefore 
take the single reference of the 6tli edition : — Yespei'tilio 
americanus vulgaris, Seba, Thes. i. p. 90, t. 55. f. 2, of which the 
original specimen (B.M. no. 67.4.12.597) is the bat known until 
1892 as Hemiderma brevicaucki. 

See Thos. P. Z. S. 1892, p. 316. 

Suggested type locality : Surinam. 



OF THE TRXTH KDITIOX OF LINX.iJUS. 131 

4. SPASMA. p. 32. Megachrma. 

The Liiinean reference (Syst. Nat. (6) p. 7) refers alone to 

Glis volans ternatanus, Seba, Thes. i. p, 90, p. 56. f. 1, which 
is the bat known as 

Megadertna spasma. 

See K. Anders. & Wrought. Ann, Mag. N. H. (7) xix. p. 132, 
1907. 

5. LEPORiNUS. p. 32. Noctilio. 

Equally through Syst. Nat, (6) p, 7, the first Linnean refer- 
ence, and Seba, pi. Iv. fig. 1, we get to the correctly known 
Noctilio leporinus. 
Suggested type locality : Surinam. 

6. AURiTus. p. 32. Plecotus. 

References: Frisch, Av, (Vogel Deutsehlands) 103. 

Edw. Av. t. 201.f. 3. 

Olear. Mus. t. 15. f. 1. 
The Long-eared Bat, Plecotus auritus. 
Type locality, from Frisch : Germany (Berlin), 

7. MURtNus. p. 32. Vespertilio. 
Linnean references : Faun, Suec, 18 ; Syst. Nat. (6) p. 7, no. 2. 

For reasons explaining why this should be considered as the bat 
formerly known as Vespertilio or Vesperugo discolor, see Miller, I. c. 
Type locality : Sweden (Upsala). 

Genus 5. Elephas. p, 33. 
Monotypic. Type, E. maximus (U, indicus auctorum). 
Species : 

MAXIMUS. p. 33. Elephas. 

Linnean reference : Syst. Nat. (6) p. 11, where the single quo- 
tation is to Ray, Quadr. p. 123 (131 in the Museum copy). 
Ray gives a long account of the Elephant, but does not 
distinguish the two forms, nor give any locality. Linnfeus, 
howevei", says " Habitat in Ceylonse paludosis," whence 
Type localit}'' : Ceylon. 

Genus 6. Trkjhecuus. p. 34. 
Monotypic. Type, T. manatits. 
Species : 

MAXATUS. p. 34. Trichechus. 

Linnean reference : Syst. Nat. (6) p. 39 (misprinted 37). 
Single reference in 6th edition : Artedi, Synonym. 109 
(misprinted 107). 
Artedi here, again, gives a primary reference to his own ' Genera 

9* 



132 MK. OLBFIELB THOMAS UN THK MAMMALS 

Piscium,' no. li. p. 556, where we find a general account which 
includes both Manatees, American and African, and Dugongs. 
Linnfeus, however, puts " in mare americano " as locality, and this 
would fix the name on the common American species. 
Type locality : "West Indies. 

Genus 7. Brabypus. p. 34. 

Two species. Type by selection : B. tridactylus. Selected by 
llliger, who at the same time eliminated B. didactyliis as a 
C'holoepus, 

Species : 

1. TRIDACTYLUS. p. 34. Bradypus. 

First Linnaean reference : Syst. Nat. (6) p. 3. 
Two references : Seba, Thes. i. t. 33. f. 2, and Marcgr. Bras. 
221, of which the first may be taken as primary. 
Type locality (suggested): Surinam. 

2. DIDACTYLUS. p. 35. Cholcepus. 

First Liiniean reference : Syst. Nat. (6) p. 3. 
One refei-ence only : Seba, Thes. i. t. 34. 
Type locality (suggested) : Surinam, 

Genus 8. Myrmecophaga. p. 35. 

Three species. Type by elimination : M. trida.ctyla, M. didactyla 
having been taken out as Cyclopes and M. ' tetradactyla as 
Taniandua. 

Species : 

1, DIDACTYLA. p. 35. Cy dopes. 
Based on specimen. Fuller description Mus. Ad. Frid. p. 8. 
No locality, but as an identification is made, through Syst. 

Nat. (6) p. 8, with Seba, pi. 37. fig. 3, we may take as the 
Type locality : Surinam. 

2. TRiDACTYLA. p. 35. Myrmecophciga. 
Linnean reference : Syst. Nat. (6) p. 8, where we find " Ray 

Quadr. 241. Marcgr. bras. 225." 

Ray's account simply refers to, and is based on, that given by 
Marcgrave of the Tamandua guacu, the Great Anteater, of which 
he gives a recognizable figure. 

The Great Anteater, commonly known as Myrmecophaga juhata, 
should therefore, as pointed out in 1901, be known as M. tri- 
dactyla. 

Type locality : Pernambuco. 

See Thos. American Naturalist, xxxv. p. 143, 1901. 



OF THE TENTH EDITION OF LIXN.5EUS. 133 

3. TETRADACTYLA. p. 35. Tamaiidua, 

Linnean reference : Syst. Nat. (6) p. 8, which gives " Ray, 
quadr. 242. Marcgr. bras. 226." 
Here, again, as in the last species, Ray simply refers back to 
Marcgrave, whose animal is the Tamandua, 

Hence the common name Tamandaa tetradactyla is correct. 
Type locality : Pernambuco. 

Genus 9. Manis. p. 36, 
Monotypic. Type, M. pentadactyla. 

Species : 

PENTADACTYLA. p. 36. . Manis. 

Linnean reference : Syst. ISTat. (6) p. 8, where four quotations 

are given : — 
Bontius, Ind. 60 (1658). 
Clus. exot. 347. 
Clear. Mus. 8, t. 7. f. 1. 
Ray, Quadr. 247. 
Of these, Bontius, as being the first quoted, should be taken ; 
and here we find the actual locality mentioned, namely, Tajoan = 
Tai- wan = Formosa. 

On this account the species which has ordinarily borne the 
name of M. dalmani Sund., should be considered as the true 
M. pentadactyla of Linnaeus. 

Moreover, Linnaeus also includes in his sjmonymy Dalman's 
paper (Act. Stockh. 1749, ]). 265) on a Pangolin from China or 
Formosa ; and as he must have seen the specimen when he gave 
the name pentadactyki, the propriety of the above identification 
is confirmed. 

Type locality : Formosa. 

Genus 10. Phoca. p. 37. 

Four species. Type by tautonymy : Ph. vitulina (" Phoca seu 
Vitulus marinus," Gesner). 

Species : 

1. URSiNA. p. 37. Callorhinus. 
Sole reference : Steller, Nov. Comm. Petrop. ii. p. 331 (1751). 

The Fur-Seal, commonly known as Callorhinus ursinus. 
Type locality : Bering Island. 

2. LEONiNA. p. 37. Mirounga. 

Sole reference : Anson, Itin. p. 100, 

Sea-Lion, 

Type locality : Juan Fernandez. 



134 MR. OLDFIELD THOMAS OX THE MAMMALS 

3. ROSMARUS. p. 38. Odobemcs. 
Linnean quotation : Syst. Nat. (6) p. 6, no. 2 (misprinted 8). 

Oori-ectly known as Odoheiius rosmarus (not OdohcBmis, if 
Brisson is taken as the founder of the generic name). 
Type locality : Northern Atlantic. 

4. viTULiNA. p. 38. Phoca. 
Linnean reference: Syst. Nat. (6) p. 6, no. 1, which refers 

again to Fauna Suecica, no. 1 1 (p. 4). 
Common Seal. 
Type locality : " in Mari Bothnico et Baltico." 

Genus 11. Canis, p. 38. 

Seven species. Type by tautonymy : Canis familiaris, among 
whose synonyms there appears " Canis Gesner." 

Species : 

1. familiaris. p. 38. Cauls. 
Linnean references : Faun. Suec. no. 12 ; Syst. Nat. (6) p. 5. 

Type locality : Sweden (Upsala). 

2. LUPUS, p. 39. Canis. 
Linnean references : Faun. Suec. no. 13 ; Syst. Nat. (6) p. 5. 

Type locality : Sweden. 

3. HY^NA. p. 40. Ilycena. 

Linnean reference : Syst. Nat. (6) p. 5, where the single 
reference is to Ksempf, Amcen. p. 407, fig. 4 (letterpress 
on p. 411). 
The animal commonly called Hycena striata, but whose proper 
name is Hymia hycena. 

Type locality: Benna Mts., Laristan, S. Persia, Kjempfer 
also mentions a sjoecimen fi^om Isfahan. 

4. vuLPES. p. 40. Vulpes. 
Linnean reference : Syst. Nat. (6) p. 5, no. 6, 

Coi-rect name : Vidpes vidpes. 
Type locality : Sweden (Upsala). 

T). ALOPEX. p. 40. {V'ldjjes.) 

Lipnean i-eference : Syst, Nat. (6) p. 5, no. 5, where the 

whole entry is 
" Oanis Cauda recta extremitate nigra Gmel. Yulpes cam- 
pestris." 
I cannot identify this animal. It has been supposed to be an 
individual variety of V. vnlpes, Avhich Blanford (P. Z. S. 1887, 
p. 635) proposed should bear the name I^, atopex. 



OF Till': L'KXTII EDITION OF LTXK^EUS. 135 

6. LAGorus. p. 40. ' Alo2)ex. 

Linnean reference : Syst. ISTat. (6) p. 5, where V. alba and 
V. ccerulescens are quoted from Kalm and the ' Fauna 
Suecica.' 

Arctic Fox. 

Type locality : Sweden (Lapland). 

7. AUREUS, p. 40. Canis. 

No Linnean reference, that to the 6th edition being omitted. 
However, the first reference, Kaempf. Amoen. p. 413, fig. 3, 
is the one quoted in that edition, p. 5, no. 5. 

Jackal . 

Type locality: Bennd Mts., Laristan, 8. Persia. 

Genus 12. Felis. p. 41. 
Seven species. Type by tautonymy ; Felis catus (" Felis" A.ldr.). 

Species : 

1. LEO. p. 41, Felis. 

Linnean reference: Syst. Nat. (6) p. 4, no. 1, where the 
quotation runs "Leo. Dod. gall. t. 1 & 17." The plate 1 
referred to appears to be that in Mem. Acad. Sci. Paris, iii. 
1666-1699, p. 1, pi. i., publ. 1733, where there is an account 
of the dissection of four lions and three lionesses. 
No type locality available. 

2. TIGRIS, p. 41. Felis. 

Linnean reference : Syst. Nat. (6) p, 4, no, 2, where two 

quotations are given : — 
Ray, Quadr, p. 165; Bont. Jav. p. 53. 
Taking the first of these, we find a description which mixes up 
the Jaguar with the Tiger, but Linne's " Habitat in Asia " may 
be taken as restricting the name to the Tiger. 

Later revisers of the races of Tigers, of whom Fitzinger and 
Matschie may be specially quoted, have restricted F. tigris to 
India proper, and particularly Bengal. 

3. PAROUS, p. 41. Felis. 

Linnean reference : Syst. Nat. (6) p. 4, no. 3, whei-e we find 
Ray, Quadi\ p. 166, and Alp. Aegypt. p. 237. 
Ray gives no indication of locality, but passing to Alpinus we 
find an account of Leopards seen at Cairo and Alexandria alive in 
captivity. The account is, no doubt, partly based on Hunting 
Leopards (Ci/nceliit'ios), but, none the less, may be accepted as 
giving for the 

Type locality : Egypt. 



J 36 MK. OLDFIELD THOMAS ON THE MAMMALS 

It may be here reiterated that Linne's " in Indiis " cannot be 
taken to mean India in the modern sense, but means hardly more 
than " from abroad." In this case, he gives a synonymy based on 
African and Mexican references, and then simply puts "Habitat in 
Indiis." 

4. ONCA. p. 42. Felis. 

First reference: Ray, Quadr. p. 168; and this in turn is 
based primarily on Marcgi'ave's account of the Jaguar. 
Consequently 
Type locality : Pernambuco, 

5. PARDALis. p. 42. Felis. 

Linnean reference : Syst. Nat. (6) p. 4, no. 4, where the only 

citation is 
" Catus pardus, Hernandez, Mex. p. 512," 
Ocelot. 

Type locality : Mexico, 

Linnasus also saw and described as the same species a living 
specimen " in Vivario cliffortiano" but there is nothing in this 
description inconsistent with the taking of the Mexican Ocelot as 
the typical form, 

6. CATus. p. 42. Felis. 

Linnea,n references : Faun. Suecica, no. 3 ; Syst. Nat. (6) 
p. 4, no. 6. 
Clearly the " Blotched Tabby " Domestic Cat., 
See Pocock, P. Z. S. 1907, p. 149, 
Type locality : XTpsala, 

7. LYNX. p. 43. Lynx. 

Linnean references: Faun. Suecica, no, 4; Syst, Nat. (6) 
p. 4, no. 7 ; It. Wgoth, p. 222. 
North European Lynx. 

Type locality : Wennersborg, S. Sweden. (From the first 
mention in the Westgotha Resa.) 

Genus 13. Viveera. p. 43. 
Five species. 
Type, both by elimination and selection : V . zibetha. 

Species : 

1. ICHNEUMON, p. 43. Mungos. 

Linnean references: Linn, in Plasselq. Itin. p. 191. 1757, and 
Syst, Nat. (6) p. 6, no, 3. 
Common Mongoose. 
Type locality : Egypt, 



OF THE TKXTU EJ^ITIOX OK LIXX.EUS. 137 

2. MEMPHITIS. p. 44. C'^) 

Iso Linnean reference. 

" Y.sqiiiepatl, Hern. Mex. 332; Seb. Mus. i. p. 68, pi. xlii. 

fig. 1 ; Rai, quadr. p. 181." 

It does not seem possible to determine to what animal this 

name should be applied. Hernandez gives a figure which has a 

superficial resemblance to a Ta,yra, but the description of the 

tail is quite inconsistent with this idea, and is applicable rather 

to a Skunk or Bassariscus. Seba's animal was probably a JVasua. 

Linnaeus himself says in one place that the colour is brown 

above and yellowish below, and in another that it is white above 

and vai-iegated black and white below, an inconsistency which 

justifies one in treating the name as unrecognizable. 

3. PUTORius. p. 44. Spilogale. 

Ko Linnean reference. 

" Putorius americanus striatus, Catesb. Carol, ii. p. 62." 
Alleghenian Spotted Skunk. 
Type locality : South Carolina. 
See Howell,' N. Am. Faun. no. 26, pp. 15-17, 1906. 

4, ziBETHA. p, 44. Viverra. 
No Linnean references, that in the 6th edition, p. 6 (Meles, 

no. 2), being omitted in the 10th. It quotes " Ray, Quadr. 
178; Dod. gall. 155." 
General references to Clusius, Ray, Dodart, Hernandez, 
Olearius, and Beltonius. 
From the name zihetha we may take the " Animal zibethicum" 
of Ray, Dodart, Hernandez, and Olearius as the primary reference, 
and of these we should take the first in Ray, who is also the first 
quoted in the omitted 6th edition reference. 

Turning to Ray we find he quotes the description by Faber in 
Hernandez, Nov. Hispan. pp. 538, 580, and 581 (also quoted by 
Linnaeus). Taking, again, the first of these we find an animal de- 
scribed which has a ringed ta.il, and would therefore be an Indian 
Civet. The illustration does not show the rings properly, but 
they are sufficiently indicated in the description. All the other 
references are to the African V. civetta. 
Type locality (fi'om Faber) : Bengal. 

5, 6EXETTA. p. 45. Geneita. 

No Linnean reference, 

"Genetta s. Ginetta, Rai, quadr, 201. Bell. itin. 76. Gesn. 
quadr. 550." 
The first reference in Ray being to Spain, that may be taken as 
the type locality, especially as the basis of the name ((lineta) 
remains as the Spanish colloquial term for the animal, 



138 MR. OLDFIELD THOMAS ON THK MAMMALS 

Genus 14. Mustela. p. 45. 

Nine species. 

Type by tautonymy : M. erminea ("Mustela" Gesner ; mis- 
quoted Mustela vulgaris), 

Althovigh in this instance alone of the Gesner quotations of the 
' Systema ' a second name is attached to the piimaiy one, for which 
tautonymy is claimed, this proves to be a misquotation by Linnseus, 
and the correct quotations seem actually to strengthen the case 
for the Ermine being really the typical tautonymous species. For 
though Linnfeus quotes " Mustela vulgaris, Gesner," we find, as a 
matter of fact, that Gesner* did not -put Mustela vulgaris, but 
simply "Mustela" in the German edition (that quoted in the 
1758 ' Systema ') and in the Latin one " MiLstela proprie sic dicta," 
in contradistinction to his " Mustelis diversis," which included the 
Marten and Polecat. 

The Ermine was therefore clearly the Mustela of both Gesner 
and Linnseus, and should be treated as the type species. 

Species : 

1. LUTEis. p. 45. Latax. 

No Linnean reference. 

Lutra marina, Act. Petrop. 1749, p. 267, i. e. Steller, Nov. 

Comm. Petrop. ii. p. 367 (1751). 
Lutra brasiliensis, Rai, quadr. p. " 187 " (189). 
The primary reference is clearly to Steller, and the type 
locality Kamtchatka. 

2. lutra. p. 45. Latra. 

Linnean reference: Faun. Suecica, no. 10. 

European Otter. 
Type locality : TJpsala. 

3. GULO. p. 45. Gulo. 

Linnean reference : Faun. Suecica, no, 6, 

Glutton. 

" Habitat in alpibus Lapponicis, Dalekarlicis, sylvisque 
vastissimis." 

Taking the first place mentioned we have 
Type locality : Lapland. 

4. BARBARA, p. 46, Tayra. 

Tayra, by description. 

" Habitat in Brasilia, M. Ac. Holmens." 

* The pagination of Gesner quoted by Linnaeus is somewhat confusing. While, in 
the majority of cases (e. (f., the Puforms), the pages are quoted from the 1620 Latin 
edition, the " Martes " and the '' Mustela vulgaris " are given as " 151 " and " 149," 
these pages coming from the German edition of 1606. Their respective numbers in 
the 2nd Latin edition (1620) are 764 and 753, and in the original edition (1551) 865 
and 851. 



■ OF THE TENTH EDITION OF LINN.EUS. 139 

5. MARTES. p. 4G. Maries, 

Linnean references : Faun. Suecica, no. " 3" (7); >Syst. Nat. 
(6) p. 5, no. 2. 
Pine Marten. 
Type locality : Upsala. 

6. PUTORius. p. 46. Mitstela. 

Linnean references : Faun. Suecica, no. 8 ; Syst. Nat. (6) 
p. 5, no. 3. 
Polecat. 
Type locality : Scania,, S. Sweden. 

7. FURO. p. 46. Mustela. 

No Linnean reference. 

" Mustela sylvestris, Viverra dicta, Rai, quadr. 199." 
Ferret. 
" Said to be from Africa." 

8. ziBELLiNA. p. 46. Maries. 

No Linnean refei-ence. 

"Mustela s(z)ibellina, Rai, quadr. p. 201." 
Sable. 
Type locality : N. Asia. 

9. ERMiNEA. p. 46. Mustela. 

No Linnean reference. 

" Mustela vulgaris, Gesn. quadr. 149 " (German edition). 
Stoat or Ermine. 
Type locality ; " In unseren Landen " (Germany). 

Genus 15. Ursus. p. 47. 

Four species. Type by tautonymy : 6''. arctos (" Ursus, 
Gesner"). 

Species : 

1. ARCTOS. p. 47. Ursus. 

Linnean I'eferences : Faun. Suecica, no. 2 ; Syst. Nat. (6) 
p. 4, no. 1. 
European Brown Bear. 
Type locality (ex Faun. Suec.) : Nortbein Sweden. 

2. Luscus. p. 47. Gido. 
No Linnean reference. 

TJrsulus lupo affinis americanus, Edw. av. ii. p. 103, t. 103, 
1747. 
American Glutton. 
Type locality : Hudson's Bay. 



140 MK. OLDFTELU THOMAS ON THE MAMMALS 

3. MELES. p. 48. Meles. 
Linnean references : Faun. Suec. no. 15 ; Syst. Nat. (6) p. 6. 

European Badger. 
Type locality : Upsala. 

4. LOTOR. p. 48. Procyon. 
Linnean references: Syst. Nat. (6) p. 4 ; Act. Stockh. 1747, 

t. 9. f. 1 (p. 277 & seqq.). 

Although in the last-named place Linufeus refers to earlier 
accounts, yec his description of an actual specimen, whose habit, 
as mentioned by him (p. 282), of dipping its food in water would 
have suggested the name lotor, should ovei'ride his synonymy. 
His statement that the Raccoon is called Ispa.n by the Swedes 
in Pennsylvania may be accepted as defining the locality of his 
animal. Therefoi-e 

Type locality : Pennsylvania. 

Genus 16. Sus. p. 49. 
Four species. Type by tautonymy : Sus scrofa ('' Sus, Gesn."). 

Species : 

1. SCROFA. p. 49. Sus. 

Linnean references : Faun. Suec. no. 36 ; Syst. Nat. (6) p. 12. 

European Wild Swine. 

Type locality : Geriiiany, whence the wild specimens referred to 
by Linnaeus were introduced * into the island of Oeland. 

2. POROUS, p. 50. Foiamochcerus. 
Linnean reference : Syst. Nat. (6) p. 12. 

Refers to " Rai, quadi-, 96," where a further reference is given 
to Marcgrave's Porcus guineensis, whose description is clearly 
that of the West- African River-Hog. 

Type locality : Guinea. 

3. TAJACU. p. 50. Tayassu. 
Linnean reference : Syst. Nat. (6) p. 12, no. 3. 

Here the only quotation is Ray (Quadr. p. 97), who in turn 
gives primary importance to Tyson's description (Phil. Trans. Roy. 
Soc. xiii. p. 359, 1683) of what he calls a Mexican Musk-Hog. 
We may therefore consider the 

Type locality : Mexico. 

4. BABYRUSSA. p. 50. Bahirussa. 

Linnean reference : Syst. Nat. (6) p. 12, no. 4, whei-e we find 
two quotations : Ray, Quadr, p. 96, and Seba, Thes. i. p. 80, 
pi. 1. figs. 2 & 3. 

Babirusa. 

Type locality (from both Ray and Seba) '. " Island of Boero." 

* See Lilljehovg, Sver. U.vgg. D;iggrlj. pt, ii, p, 761, 1874. 



OF THE TICNTH KDrrU)X OF I.lXX.tlS. 141 

Genus 17. Dasypus. p. 50. 

Six species. Type by tautonymy: /). novemcinctns (•'Dasypu.s, 
Hern."). 

Apart from the difficult and doubtful ca,se of Simla, this is the 
only instance in which the use of tautonymy in selecting Linnean 
types brings out a result conti'ary to common use. 

Linnteus placed the name " Dasypus'' among the synonyms of 
D. novemcinctus alone, and the account in Hernandez from which 
this name comes is cleai'ly that of one of the group long known as 
Tatasia, more recently as Tatu. 

The comparatively recent date of the latter cha.nge, the extreme 
ugliness of I'atu, which antedates 7'atusia. and the classical suita- 
bility of Eaphractas, which stands for sexcinctits and its allies, 
will all help to i-econcile us to the shifting of Dasypios from one 
group to another. 

This shifting is quite unavoidable, if the invaluable principle of 
tautonymy is to be utilized at all- 
Species : 

1. UNiciNCTUS, p. 50. Cabassous. 

Linnean reference : Syst. ISTat. (6) p. Q, no. 4, where the sole 
reference is to Seba, Mus. i. p. 47, pi. xxx. figs. 3 & 4. 
This, or at least fig. 3 (fig. 4 being more doubtful), is clearly the 
Ta,touay or Twelve-ba.nded Armadillo. 

No original locality available., but being' from Seba, Surinam 
may be suggested, if Cabassous occurs there, which is not as yet 
certainly known. 

2. TRiomcTus. p, 51. ' Tolypeutes. 

Linnean reference : Syst. Nat. (6) p. 6, no. 2, where the sole 
quotation is to Seba, i. p. 62, pi. xxxviii, figs. 2-3. 
Seba's animal is a typical Three-banded Armadillo- 
Suggested type locality : Surinam. 

3. QUADRiciNCTUS. p. 51. Tolypeutes. 

Linnean reference : Syst- Nat. (6) p. 6, no. 3, where the single 
quotation is to "Column, aquat. ii. p. 15. pi. xvi." (1606). 
Oolonna's animal is a four-banded specimen of Tolypeutes 
tricinctus, of which D. quadricinctus is ther-efore a s\'nonym. 

4- SEXCINCTUS. p. 51. Euphractus. 

Linnean references: Mus. Ad. Frid. p. 7 (1754), and Syst. 
Nat. (6) p. 6, no. 5. 

The former takes precedence, as being based on a specimen, 
whatever the reference to Ray in the latter might lead to. 

Six-banded Armadillo. 

Type specimen in Stockholm Museum, its locality unknown, 
Tint Para specimens agree with it in size, and ma v be provisionally 
accepted as typical {cf. P. Z. S. 1903, ii. p. 242 j. ' 



142 MR. OLDFIELB THOMAS ON THE MAMMALS 

5. SEPTEMCiNCTUS. p. 51. J)as i/jjus {= Tcitu axict.). 

Linnean references: Amcen. Acad. i. p. 281, and Syst. ISTat. 
(6) p. 6, no. 6 ; but in the latter we are again referred to 
the former. Thus the synonymy may be excluded and 
primary importance given to the species described. This 
is said to be still in the Upsala Museum. 

Seven-banded Armadillo. 

Type locality not known. 

6. NOVEMCINCTUS. p, 51. Dasypus. 
Linnean references : Mus. Ad. Frid. p. 6, and Syst. Nat. (6) 

p. 6, no. 7, but in the latter no further detail is added 
and we must take the former alone. 
Nine-banded Armadillo. 

Prof. Einar Lonnberg informs me that the type specimen is still 
preserved in the Stockholm Museum, and this will need expert 
examination whenever an attempt is made to woi-k out the 
present difficult group. 

Genus 18. Erinaceus, p. 52. 

Monotypic. Type, E. eim-opceus. 

Species : 

EUROP^us. p. 52. Erinaceiis. 

Linnean I'eferences to Fauna Suecica, Syst. Nat. (6), and 
Iter Goth, 
Common Hedgehog. 

Tj'pe locality (from the Gothlandska Resa, p. 264) : Wamlingbo, 
S. Gothland Id. 

Genus 19. Talpa. p. 52. 
Two species. Type by tautonymy : T. europcea (" Talpa, Gesn."). 

Species : 

1. europ^a. p. 52. Talpa. 
Linnean references to Syst. Nat. (6), Fauna Suecica, and 

Iter Scanicum. 
Common Mole. 
Type locality : Upsala *. 

2. ASiATiCA. p. 53. Chrysochloris. 
Sole reference : Seba, Mus, i. p. 51, pi. xxxii. figs. 4, 5. 

Cape Golden Mole. 

Type specimen in British Museum (Lidth de Jeude Coll.), 
B.M. No. 67.4.12.564 (see P.Z.S. 1892, p. 316). 

* The ' SkSnska Resa ' being later in date than the ' Fauna Suecica,' one cannot 
justifiablj' fix a type localitj' from it wlicn the ordinary Ups^alan locality is available 
from the latter work. 



OF TUK TKXTil KDITIOX OF LIXX.F.IS. 14i5 

Genus 20. Sorex. p. 53. 

Thi-ee species. Type by tautonymy : S. aranens (" Sorex, Faun. 
Snec."'). 

tSpecies : 

1. ARANEUS. p. 53. Sorex. 
Linnean references: Faim. Suec. no. 33 ; Syst. Nat. ((5) p. 10. 

Common Shrew. 
Type locality : Upsala. 

2. CRiSTATUS. p. 53. Comlylara. 
Sole reference : P. Kalm. 

Type locality : Pennsylvania. 

3. AQUATicus. p. 53. Scalops. 

Reference to Seba (pi. xxxii. fig. 3), whose figure is of Talva 
europcea, but the description is cleai-ly that of an American 
Mole, and was no doubt received from Kalm, who is quoted 
as authority for the locality. 

Common Ameiicaii Mole. 

Type locality : Philadelphia. 

Genus 21. Didelphis. p. 54. 
Five species. Type by selection and elimination : D. marsupialis. 

Species : 

1. MARSUPiALis. p. 54. Didelphis. 

Linnean references: Auicen. Acad. i. p. 279; Syst. Nat (6) 
p. 10; Mus. Ad. Fr. p. 10. 
In the first of these places the primary, and in the others the 
sole reference is to Seba, Mus. i. p. 64, pi. xxxix., which should 
be taken as the basis of the name. 
Suggested type locality : Surinam. 

2. PHILANDER, p. 54. Caluromijs. 
Sole reference : Seba, Mus. i. p. 57, pi. xxxvi. fig. 4. 

Philander Opossum. 

Type specimen in British Museum, No. 67.4.12.414 (see P.Z.S 
1892, p. 314). 

Suggested type locality : Surinam. 

3. OPOSSUM, p. 55. Mefackirus. 
Sole reference: Seba, Mus. i. p. 56, pi. xxxvi. figs. 1-2. 

Quica Opossum. 

Suggested type locality : Surinam. 



]44 MR. OLDKIKIiL) THOMAS ON THE MAMMALS 

4. MURINA. p. 55. Marmosa. 

Linneaii reference : Amoen. Acad. i. p. 279. — This is an 
error, the only Opossum referred to being the large 
Dide IjjMs m arsupicdis . 
Second reference : Seba, Mus. i. p. 48, pi. xxxi. figs. 1-2, 
which must be taken as the ba.sis of the name. 
Murine Opossum. 

Co-types in British Museum, Nos. 67.4.12.541-542 (see P. Z. S. 
1892, p. 314). 

Suggested type locality : Surinam. 

5. DORSiGBBA. p. 55. Mcirmosa. 
Primary reference to Seba, Mus. i. p. 49, pi. xxxi. fig. 5. 

Synonym of Marmosa murina. 

Type in British Museum, No. 67.4.12,546. 

Suggested type locality : Surinam. 

Oenus 22, Rhinoceros, p. 56. 

Two species. Type by tautonymy : R. unicornis ('• Rhinoceros, 
Jonst."). 

Species : 

1. UNICORNIS, p. 56. ■ Rhinoceros. 
Linnean reference : Syst. Nat. (6) p, 11, no, 1. 

One-horned Indian Rhinoceros. 

From Jonston's 'Quadrupeds' (p. 67, 1657), the first work 
quoted in the 10th edition, we get the 
Type locality : Bengal. 

2. BicoRNis. p. 56, Diceros. 
Linnean reference: Syst. Nat, (6) p, 11, no. 2, where no 

further quotation is given. 

Always accepted as the African Two-horned Rhinoceros. 

Type locality not determinable, except that the Cape may 
generally be considered the place whence early specimens wei'e- 
brought '' Habitat in India " may, as usual, be entirely ignored. 

Genus 23. Hystrix. p. 56. 
Five species, T3'pe by tautonymy : R. crisfate(" Hystrix, Gesn."). 

Species : 

1. CRIST ATA. p. 56. Hystrix. 

Linnean reference : Syst. Nat. (6) p. 9, no. 1, where the fir,st 
quotation is to Ray (Quadr, p. 2U6), whence we find 
Type locality : Ronae. 
Common Porcupine, 



\ OF THE TE.STH EDITIuN OF LINN.EC3. 145 

2. PREHENSiLis. p. 57. Coendou. 

Linnean reference : Syst. ]S"at. ((5) p. 9, no. 2, where the only 
quotation is to E,ay (Quaih-. p. 208). There we get an 
account taken from Mavcgi-ave, whence 

Type loeahty : Pernambuco. 

Brnzilian Poi'cupine. 

3. DORSATA. p. 57. Erethizon. 

Sole reference : Edw. Aves, p. 52, pi. lii. 

Oanadiai^ Porcupine. 

Type locality : Hudson's Bay. 

4. MACROURA. p. 57. Atherurvs. 

Linnean reference: Syst. Nat. (6) p. 2, no. 3, where the sole 
quotation is Seba, Thes. i. p. 84, pi. lii. fig. 1. 
Common Brush-tailed Porcupine. 
No type locality available. 

5. BRACHYURA. p. 57. . Acanthxon. 

Linnean reference : Syst. Nat. (6) p. 9, no. 4, where the sole 
quotation is Seba, Thes. i. p. 81, pi. li. fig. 1. 
Malay Porcupine. 
Type locality : Malacca. 
See Lyon, P. U.S. Nat. Mus. xxxii. p. 579, 1907. 

Genus 24. Lepus. p. 57. 
Four species. Tj'^pe by tautonymy: L. timidus (" Lepus, Gesn."). 

Species : 

1. TIMIDUS. p. 57. Lepus. 

Linnean references : Syst. Nat. (6) p. 9, no. 2 ; Faun, Suec. 
no. 19. 
"Variable Hare. 
Type locality : Sweden (Upsala). 

2. CUNICULUS. p. 58. Oryctolagus. 
Linnean references : Syst. Nat. (6) p. 9, no. 3 ; Faun. Suec. 

no. 20. 
In the former a reference is again given to the latter. 
Clearly based on the Domestic Rabbit ('• pupillis rubris"). 
Later on Linnseus became acquainted with the Wild Rabbit, 
and stated in the 10th edition : — 
" Habitat in Europa australi" 

3. CAPENSis. p. 58. Lepus. 

Common Cape Hare. 
Type locality : Cap3 of Good Hope. 

I have considered the quesbion as to whether this might not be 
Proc. Zool, Soc— 1911, No. X, 10 



146 MR. OLDFIELD THOMAS OX THE MAMMALS 

a Rooi-haas {Pronolagus), but think the length of the tail too 
great, while in any case the evidence is insufficient to vipset the 
usually accepted determination. 

4. BRASiLiENSis. p. 58. Sylvilagus. 

Linnean reference: Syst. N'at. (6) p. 9, no. 1, where the sole 
quotation is Ray, p. 205, whose account is taken from 
Marcgrave. 
Hence 
Type locality : Pernambuco. 

Genus 25. Castor, p. 58. 

Two species. Type by tautonymyj C. fiber (" Castor, Gesn.").' 

Species : 

1. FIBER, p. 58. Castor. 
Linnean references : Faun. Suec. no. 23 ; Syst, Nat. (6) p. 10, 

no. i. 
Ill the latter the only reference is to the former, in which we find 
Type locality : Lapland. 

2. MOSCHATUS. p. 59. Desmana. 
Linnean references : Faun, Suec. no. 24 ; Syst. Nat. (6) p. 10, 

no. 2. 
Type locality : Russia. 

Genus 26. Mus. p. 59. 

Sixteen species. Type by tautonymy : M. musculus (" Mus, 
Gesn."). 

Species : 

\. PORCELLUS. p. o9. Cavia. 

Linnean references : Westgoth. Resa, p. 244 (misprinted 
224); Syst. Nat. (6) p. 10, no. 1. 
Domesticated Cavy. 

2. LEPORiNUS. p. 59. Dasyprocta. 

Sole reference: Catesby, Carol, iii. (i. e. ii. Appendix) p. 18, 
pi, xviii. 
An indeterminable species of Dasyprocta, Tail too short and 
size too large for Myoprocta. 
No type locality available. 

3. LEMMUS. p. 59. Lemmus. 

Linnean references to Fauna Suecica and Syst. Nat. (6), as 
well a,s to Act. Stockh. 1740, p. 320, pi. vi. figs. 4 & 5, 
where Linnseus gives an account of the Norway Lemming. 
Type locality : La|>land, 



OF THE TEXTH EDITION OF MXX.WUS. 147 

4. MARMOTA. p. 60. " Mannota. 

Liunean reference: Sy.st. ]S"at. (6) p. 10, no. 11, wliere the 
sole (] notation is Ray, p. 221. 
Alpine Marmot. 
Type locality ; Swiss Alps. 

5. MOXAX. p. 60. Marmota. 
Sole reference: Edwards, AA^es, p. 104, pi. cv. 

Wooileliuck. 

Type locality : Maryland. 

6. cracETiJS. p. 60. Criceius. 

Linnean reference: Syst. N'at. (6) p. 10, no. 10, where the 
sole quotation is Ray, p. 221. 
" In Turingia and Misnia reperitur" — consequently 
Type locality : Thiiringen. 
Common Hamster. 

7. TERKESTHis. p. 61. Arvicola. 

Linnean references : Faun. Suec. no. 29 ; Syst. Na,t. (6) 
p. 10, no. 5. 
Swedish Water- Yole. 
Type locality : Upsala. 

8. AMrHiBUJS. p. 61. Arvicola. 
Primaiy reference : Ray, Quadr. p. 217. 

Ray's account is quoted from Willughby, and, as both were 
Englishmen, the animal should be taken as the British Water- 
Yole. 

Type locality : England. 

9. RATTUS. p. 61. Epimys. 

Linnean references: Fnun. Suec. no. 28 ; Syst. Nat. (6) p. 10, 
no. 6. 
Black Rat. 
'IVpe locality : Sweden (Upsala). 

10. MusctTH's. p. 62. Mus. 

Ijinne;ui references : Faun.Suec.no. 31 ; Syst. Nat. (6) p. 10, 
no. 8. 

House- Mouse. 

Type locality : Sweden (Upsala). 

11. AVELLAXARirs. p. 62. Muscardinus. 
Linnean reference : F^ann. Suec. no. 32. 

]~)ormouse. 

Type locality : Swedt^i (T'psala). 

JO* 



148 MR. OLDFIELD THOMAS O.N THE MAMMALS 

12. SYLVATicus. p. 62. Apodemus. 
Linnean references : Faun. Suec. no. 30 ; Syst. Nat. (6) p. 10, 

no. 7. 
Long-tailed Field-Mouse. 
Type locality : Sweden (Upsala). 

13. STRiATUS. p. 62. Arvlcmithis. 
Linnean reference : Mus. Ad. Frid. p. 10. 

The punctated white lines described indicate that this mouse 
is the species usually known as Arvicanthis pulchellus Grny. 
■Having been brought in a ship coming from " India," its locality 
is no doubt that part of W. Africa where such a ship would ha\ e 
touched. Consequently we may consider its 

Type locality : Sierra Leone. 

14. LONGiPES. p. 62. Allactaga. 

Linnean reference : Mus. Ad. Frid. p. 9. 

A five-toed Jerboa, not specifically determinable without 
examination of the type, if one exists. 

15. JACULUS. p. 63. Jaciilus. 

References : Linn, in Hasselq. Itin. p. 198 ; Act. Stockh. 
1752, p. 123. 
Lesser Egyptian Jerboa.. 
Type locality : Lower Egypt. 

16. voLANS. p. 63. Sciuropterus. 
Primary reference: Ray, Quadr. p. 215, whence we get 

Type locality : Virginia. 

American Flying-Squirrel. 

Other references are given to Seba, who figures a young 
Petaurista, and Edwards, whose animal was drawn from a living 
specimen brought from N. America. 



Genus 27. Sciurus. p. 63. 

Seven species. Type by tautonymy : S. vulgaris ("Sciurus, 
Gesn."). 

Species : 

1. VULGARIS, p. 63. Sciurus. 

Linnean references: Faun. Suec. no. 2l ; Syst. Nat. (6) p. 9, 
no. 1. 
Common Squirrel. 
Type locality : Sweden (Upsala), 



OF THE TEXTM EDITION' OF LINN.T^US. 149 

2. NIGER. p. 64. 

Sole reference : Catesbv, Carolina, ii. pi. Ixxiii. 
Southern Fox-Squirrel. 

Type locality : Carolina. Considered by Bangs to be Southern 
South Carolina. 

See Bangs, P. Biol. Soc. Wash. x. p. 147, 1896. 

3. ciXEREus. p. G4. Sciurus. 

References: Riy, Quadr. p. 215; Catesby, Carolina, ii. p. 74, 
pi. Ixxiv. ; Kalin. Itin. ii. p. 409. 
Ray's account is short and unimportant, and Catesby's is clearly 
the primary one. Mr. Bangs has allocated Catesby's animal to 
the Southern Fox-Squirrel (Scmrios niger) in his work on Eastern 
N.- American Squiri'cls, and his determination may be accepted. 
Type locality : coast region of Carolina. 

4. FLAVUS. p. 64. 

Linnean and sole reference: Amoen. Acad. i. p. 281. 
*' Habita,t in America." 

Indeterminable, except by reference to the type, if one exists. 
In the 12th edition, Carthagena is added as the locality, but 
on wliat gi'ounds does not appear. 

5. GETULUS. p. 64. Atlavioxerus. 

References; Ray, Quadr. p. 216; Edwards, Aves, pi. 198; 
Seba, Mus. i. p. 76, pi. xlvii. fig. 3. 
The first two both refer to the Barbary Squirrel, and the third, 
which is a Chipmunk, may be ignored. 
Type locality : Barbary. 

6. STRiATUS. p. 64. Tcomias. 

Linnean reference ; Mus. Ad. Frid. p. 8, where a quotation 
is given of Catesby, Carolina, ii. pi. Ixxv. 
Common Eastern Chipmunk. 
Type locality : " Virginia and Carolina." 

7. voLANS. p. 64. Sciuropterus. 
Linnean references: Faun. Suec. no. 22 ; Syst. Nat. (6) p. 9. 

Northern Flying Squirrel, Schiropterus russicus, the name 
volans being already used (from Mus volans L.) in the genus. 
Type locality : Finland. 

Genus 28. Camelus. p. 65. 

Four species, of which two, C. ch'omedaritis and hactrlanus, 
have " Camelus " among their synonyms. 

Type by tautonyiny and elimination : G. hactrlanus *. 

* Cf. ' Otiiiiit)iis ot Nnmem'l.Ttmx (.'ommissioners," 1010, p. 37. 



150 MR. OLDFIELD THOMAS OX THE MAMMALS 

Species : 

1. DROMEDARiTis. p. 65. Ccimelus. 

Linnean reference: Syst. Nat. (6) p. 13, no. 1, where the 
sole quotation is of Ra}', p. 143. 
One-hnmped Camel. 
Type locality : " deserts of Libya and Arabia " (Rfjy). 

2. BACTRTANUs. p. 65. Camelus. 
Linnean reference: Syst. Nat. (6) p. 13, no. 2. 

Two-humped Camel. 
Type locality : " Bactria." 

3. GLAMA. p. 65. Lama. 
Linnean I'eference : Syst. Nat. (6) p. 13, no. 3, where the 

sole quotation is of Ray, p. 145. 
Llama.. 
Type locality : Peru. 

4. PACos. p. 66. jLama. 

Linnean reference: Syst. Na,t. (6) p. 13, no. 4, where Ray 
(p. 147) is again alone quoted. 
Alpaca. 
Type locality : Peru. 

Genus 29. Mosomis. p. 66. 
Monotypic. 

Species : 

1. MOSCHIFERUS. Moschus. 

Linnean reference : Syst. Nat. (6) p. 13, where Ray is alone 
quoted. 
Type locality (from Linnfeus) : " Tataria versus Chinam." 

Genus 30, Certu.s. p. 66. 

Eight species. T^-pe by tautonymy : C. flaphus (" Cervus, 
Gesn "). 

Species : 

1. CAMELOPARDAiJS. Giraffa. 

Linnean reference : Syst. Nat. (6) p. 13, no. 1, where 
references are given to Bellonius (Obs. ii. p. 119) and Ray, 

the latter of whom merely quotes the former. 
Giraffe. 
Type locality : Egypt (seen in captivity at Cairo). 



OF THE TENTH EDITION OP^ L1NN.EU3. 151 

2. ALCEs. p. 66. Alces. 

Liuneaii references: Syst. Nat. (6) p. 13, no. 2 ; Faun. Suec. 
no. 37. 
Elk. 
'IVpe locality : Sweden. 

3. ELArHUS. p. 67. Cervus. 

Linnean references : Faun. Suec. no. 38 ; Syst. Nat. (6) 
p. 13, no. 3. 
Red Deer. 
Type locality : Sweden. 

4. TARANDUS. p. 67. Rangifer. 

Linnean references ; Faun. Suec. no. 39 ; Syst. Nat. (6) 
p. 13, no. 4. 
Reindeer. 
Type locality : Lapland. 

5. DAMA. p. 67. Cervus. 

Linnean references : Faun. Suec. no. 40 ; Syst. Nat, (6) p. 13, 
no. 5. 
Fallow Deer. 
*' Habitat in vi^•al■iis Regis & Magnatum." 

6. BEZOARTious. p. 67. Blastocerus. 

References : 

Mazama, Hern. mex. p. 324. 

Cuguacu-ete, Maregr. Bras. p. 235 ; Piso, Bras. p. 98 ; Ray, 
Quadr. p. 90. 
Linnean diagnosis taken from Marcgi-ave, whose account should 
therefore be selected as the primaiy reference. 
Pampas Deer i^Cariacus campestrw auctorum). 
Type locality : Pernambuco. 
See Lydekker, ' Deer of all Lands,' p. 287. 

7. CAPRBOLUS. p. 68. Capreohis. 

Linnean references : Faun. Suec. no. 41 ; Syst. Nat. (6) 
p. 13, no. 6, 
Roe- Deer. 
Type locality : Sweden. 

8. GUINEENSIS. p. 6H. (?) 
Linnean reference : Mus. Ad. Ki'id, i. p. 12. 

Hab. Ouinea. 

Not as vet iilentified. 



1&2 MK. OLDFIELD THOMAS ON THE MAMMALS 

Genus 31. Capra. p. 68. 
Twelve species. Type by tautonymy : C.hircus (" Capra, Gesn."). 

Species : 

1. HTRCUS. p. 68. Capra. 
Linnean reference : Faun. Suec. no. 42. 

Domestic Goat. 

2. IBEX. p. 68. Capra. 

Linnean reference : Syst. Nat. (6) p. 14, no. 6. 

Reference to Ray only — p. 77 (misprinted 79). 
Swiss Ibex. 
Type locality : Swiss Alps — Valais. 

3. KTTPICAPRA. p. 68. Rupicapra. 
Linnean reference : Syst. Nat. (6) p. 14, no. 5. 

Chamois. 

Type locality : Swiss Alps. 

4. DEPRESSA. p. 69. (?) 

Linnean reference : Syst. Nat. (6) p. 14, no. 2, where a further 
reference is given to the 3rd and 4th editions of the 
'Systema,' but no additional information is there given. 
Indeterminaltle. 

5. REVERSA. p. 69. (?) 

Linnean reference: Syst. Nat. (6) p. 14, no. 3, where also 
the 3rd and 4th editions are quoted. 
Indeterminable. 

6. PYGMEA. p. 69. Neotragus. 

Linnean reference : Syst. Nat. (6) p. 14, no. 4, where Seba 
(i. p. 70, pi. xliii. fig. 3) is alone quoted. 
Royal Antelope. 
Type locality : Guinea. 

7. GAZELLA. p. 67. Onjx. 

Linnean reference : Syst. Nat. (6) p. 14, no. 8, where Ray's 
Gazella indica (p. 79) is alone quoted. 
Common Gemsbok. 
Locality not defined, but may be taken as S. Africa. 

8. CERViCAPRA. p. 69. Antilope. 

Linnean reference : Syst. Nat. (6) p. 14, no. 7, where there 
are two quotations — 

Gazella africana, Ray, Quadr. p. 79. 

Oapricerva, K^mpf. Amoen. p. 398, pi. 401, fig. 1. 
Both refer to the Indian Black-Buck. 
Type locality t India. 



OF tup: tenth EDITIOX of IJNN.EUS. 153 

9. DORCAS, p, 69. Gazella. 

Linnean reference : (6) p. 14, no. 9, where the sole quotation 
is Ray, p. 80. 
Common Gazelle. 
No definite locality. 

10. GRiMMiA. p. 70. Cephalophus. 

Linnean reference : Syst. Nat. (6) p. 14, no. 10, where the 
only quotation is to Ray's Capra sylvesiris africana 
grimmii, which again refers to " Ephem. German. An. 14, 
Obs. 57," i. e. Grimm, Misc. cur. Acad. Nat. Cur., Decas ii. 
Ann. iv. p. 131 (1686). 

Common Duiker. 

Type locality: Cape Town. 

11. MAMBRicA. p. 70. Capra. 

Linnean reference : Syst. Nat. (6) p. 14 (no. 11), where the 
primary quotation is to Ray, p. 81. 
A long eared domestic Goat. 
Type locality : Syria. 

12. AMMOJ^r. p. 70. Ovis. 

Sole reference : Gesn. Quadr. p. 155, but the diagnosis and 
" Habitat in Siberia, Gmelin," indicate another and more 
essential source of information, which may be traced to 
J. G. Gmelin, Reise durch Siberien, i. p. 368 (footnote), 
1751. 

Siberian Wild-Sheep. 

Type locality : Upper Irtisch R., Siberia. 

Genus 32. Ovis. p. 70. 
Three species. Type by tautonymy : 0. aries ('■ Ovis, Gesn."). 

Species : 

1. ARIES. p. 70. Ouis. 

Linnean references : Faun. Suec. no. 43 ; Syst. Nat. (6) 
p. 15, no. 1. 
Domestic Sheep. 
Type locality : Sweden. 

2. GUINEENSIS. p. 71. Ovis. 

Linnean reference : Syst, Nat. (6) p. 15, no. 3, where the 
reference is to Ray, who in turn quotes Marcgrave. 
Domestic Sheep of Guinea. 

3. STREPSICEROS. p. 71. Ovis. 

Linnean reference: Syst. Nat. (6) p. 15, no. 2, whence 
Bellonius, Obs. i. p. 20, 1605. 
Cretan Domestic Sheep. 



154 MR. OLDFIIiLB THOMAS OX THE MAMMALvS 

Genus 33. Bos. p. 71. 
Five .'jpecies. Type by taiik)iiyiny ; B. taurus (" Bos, Gesn."). 
Sp?cie« : 

1. TAURUS. p. 71. Bos. 

Linnean references : Fanri. Siiec. no. 44 ; Syst, Nat. (6) 
p. 15, no. i. 
Domestic Ox. 
Ty^pe locality : Sweden (Upsala). 

2. BOXASus. p. 71. Bison. 

Linnean reference: Syst. Nat. (6) p. 15, no. 2, where there 
is no further reference, so we may take that in the 10th 
edition to the Bonasus of Ray (p. 71). 

European Bison. 

No type locality available. 

3. Bisox. p. 72. Bison. 

Linnean reference: Syst. Nat. (6) p. 15, no. 3, where the 
sole quotation is the ^^ Taitrns mexicanus" of Hernandez, 
Mexico, p. 587. 
American Bison. . 

Tjrpe locality : " Mexico." 

4. BUBALis. p. 72. Buhalus, 

Linnean reference : Syst. Nat. (6) p. 15, no. 4, where a 
diagnosis, but neither further reference nor locality, is 
given. Passing to the next reference in the 10th edition, 
we get Ray (p. 72), whose account is based on the domesti- 
cated Buffalo. Therefore 
Type locality : Italy (Rome). 

5. ixDictTS. p. 72. Bos. 
Sole reference : Edwnrds, Aves, iv. p. 200 (1751). 

Indian Humped Ox. 
Type locality : E. Indies. 

Genus 34. Equus. p. 73. 

Three species. Type by tantonymy : E. caballufi ("• Equus, 
Gesn."). 

Species : 

1. CABALLUS. p. 73. ./{qniis, 

Linnean refei-euco ; Faun, Suec, no, 34. 
Domestic Morse, 



OF THE TEXra EDIIIOX UK LINX.T;US. J f);") 

2. ASI\US. p. 73. Equus. 
Linnean reference : Faun. Suec. no. 35, 

Domestic Ass. 

3. ZEBRA. p. 74. Equus. 
Linnean reference: Syst. Nat. (6) p. 11, no. 3, where a 

diagnosis, bvit no locality or further I'eference, is given. 
Passing to the next reference we have Edwards, Aves, v. 
pp. 27 & 29, pis. 222 k 223, the first of which is a Mountain 
Zebra, and the second a Qnagga. The first should be 
ta.ken as typical. 
Type locality : Ca|)e of (lood Hope. 

Genus 35. HirroroTAMus. p. 74. 

Two species. Type by tautonymy : H. anqjhihms ('• Hippo- 
potamus, Bell.'*). 

Species : 

1. AMrHiBius. p. 74. Hippopotamus. 

No Ijinnean reference, though the species occurs Svst. Nat. 
(6) p. 11. 
Hippopotainus. 
Type locality : R. Nile. 

2. TERRKSXRis. p. 74. Tapirus. 
Two references : Ta.piierete, Marcgr. Bras. p. 229 ; Ray, 

Quadr. p. 126, where Marcgrave is again quoted. 
Brazilian Tapir. 
Type locality : Pernambuco. 

Genus 36. Moxodox'. p. 75, 
Monotypic. 

Species : 

1. MONOCEROs. p. 75. Monodon. 

Linnean references : Faun. Suec. no. 263 ; Syst. Nat. (6) p. 39, 
Narwhal. 
" Habitat in mari atlantico." 

Genus 37. Bal^exa. p. 75, 

Four species. Tvpe by tautonymy : B, mysticetus ('" Balfena, 
Will."). 

Species : 

1. MYSTICETUS. p. 75. Balcena. 

Linnean reference? : Faun. Suec. no. 264 ; Syst. Nat. (6) 
p. 39, no. 1. 
In these places the species is called '' Gronlands Waltisk " and 



156 MR. OLDFIELD THOMAS ON THE MAMMALS 

" ( i!rouia.nd.swal " respectively, so that the Greenland species of 
Iliijht Whale may be at once accepted. 

Type locality ; Greenland seas. 

Mr. True, in his ' Nomenclature of the Whalebone Whales ' *, 
comes to the same conclusion, for the same reason, after two 
pages of discussion of what I venture to consider negligible 
synonyms. 

2. PHYSALUS. p. 75. Balcenoptera. 

Linnean references : Faun. Suec. no. 265 ; Syst. ISTat. (6) 
p. 39, no. 2. 

In both these places the reference is to Artedi (Gen. 77, 
Syn. 107), to which alone validity should be attached. 

On reference to Artedi, we find Ray (Syn. Pise. 9) quoted 
primarily, who in turn gives an abbreviated account of Martens's 
" Finfisch." This has been shown by Mr. True to be a Common 
Rorqual, to which, in agreement with him, I think the name 
physalus should be applied. 

Type locality : Spitzbeig'en seas. 

The result is therefore the same as Mr. True's, but the steps are 
formal, strict, and inevitable, and do not involve any balancing 
of the importance to be attached to different synonyms. 

3. BOOPS. p. 76. BcdLBJioptera. 
Linnean reference: Mus. Ad. Frid. i. p. 51 (misprinted 50). 

But this reference is wrong as to page, and the diagnosis is a 
repetition of that of the succeeding species, B. mtisculus. 1 1 may 
therefore be ignored. 

The next reference is to species iii. on p. 107 of Artedi's 
' Synonyms,' where in turn Ray's " Balsena tripinnis nares 
habens ....'"' is quoted. The latter is based on Sibbald's 
account of a whale which Mr. True shows to be the same as 
the last species, B. physalus, of which, therefore, B. hoo])s is a 
synonym. 

Type locality : Firth of Forth, Scotland. 

4. MUSCULUS. p. 76. Balcenoptera. 
Linnean reference : Syst. Nat. (6) p. 39, no. 3. 

Here we get Artedi, Syn. p. 107, whence come references to 
Ray (Pise. p. 17), and, at last, the real basis of all, Sibbald's 
' Phalainologia.' 

Sibbald's Rorqual. 

Type locality : Firth of Forth, Scotland. 

* P. U.S. Nat. Mus. xxi. p. 617, 1898, 



OF TUE TENTH EDITION OF LIXN.EUS. 157 

Genus 38. Physeter. p. 76. 

Four species nominally, but all prove to be the Sperm Whale, 
so that the genus is really monotypic. 

Species : 

1. CATODON. p. 76. Physeter. 

Linnean reference : Syst. Nat. p. 39, no. 1 (of Catodon, not 
Physeter), where there is a reference to the " Catodon 
Ustula in rostro " of Artedi(Syn. p. 108), by whom in tui'n 
Ray and Sibbald are quoted. • 

Ray's account is abbreviated from that of Sibbald, whose 
Balaena minor *, fi'om Kairston, Orkneys, is the real basis of the 
name. 

This animal has been said by some authoi'S to be the Beluga, 
and by others the Sperm Whale, and after a careful consi<leration 
oi: the question, with the kind and highly opportune assistance 
of Dr. Einar Lonnberg, I am decidedly of opinion that it was the 
latter species. 

The absence of teeth in the upper jaw is a definite character, to 
which much weight should be attached ; while the fact that females 
of the Sperm Whale go together in schools would account for the 
large number (105) stranded at Kairston, and their comparatively 
small size (24 feet). 

As a consequence, the Sperm Whale should bear the name 
of Physeter catodon L., of which the next name would be a 
synonym. 

Type locality : Kairston, Orkneys. 

2. MACROOEPHALUS. p. 76. Physeter. 

Linnean references: Faun. Suec. no. 262; Syst. ISTat. (6) 
p. 39, no. 2 (of Catodon). 
Sperm Whale {Physeter catodmi, see above). 
Type locality : JSTorwegian seas. 

3. MICROPS. p. 76. [Physeter.) 

Linnean reference : Syst. ISTat. (6) p. 39, no. 1 (of Physeter), 
where Artedi (Syn. 104, Gen. 74) is alone quoted, 

4. TURSIO. p. 77. {Physeter,) 

Linnean reference : Syst. Nat. (6) p. 39, no. 2 (of Physeter), 

where Artedi (Syn. 104, Gen. 74) is alone quoted. 

These two names refer to the mythical " High-finned Cachalot," 

whose origin no doubt lies in faulty observation of ordinary 

Cachalots, and they may be considered as synonyms of Physeter 

catodon. 

* ' Phalaiiuilogia nova.' Caput ii. p. -li. 



158 MR. OLDFIELD THOMAS OH 

Genus 39. Delphinus. p. 77. 

Thi'ee species. Type by tautoavmy : D. deljyhis (" Delphinus, 
Bell.''). 

Species : 

1. PHOc^XA. p. 77. Phoccena. 

Linnean references: Faun. Suec. no. 266 ; Syst. Nat. (6) 
p. 39, no. 1. 
Common Porpoise. 
Type locality : Sweilish seas. 

2. UELPHis. p. 77. Delphinus. 

Linnean reference : Syst. N"at. (6) p. 39, no. 2, where Artedi 
(Syn. 105) is alone quoted. 
Conniion Dolphin. 
Type locality : " European seas." 

3. OKCA. p. 77. Orcinus. 

Linnean references : Fann. Suec. no. 267 ; Syst. Nat. (^6) 
p. 39, no. 3. 
. Common Killer. 
Type locality : North Sea. 

10. Th(^ [)ake oF Bedfonl's Zoological Exploration of Eastem 
Asia. — XI n. On Mammals from the Provinces of 
Kan-su ami Sze-chwan, Western Clhina. By Oldfikld 
Thomas, F.R.S., F.Z.S. 

[Received and Read February 7, 1911.] 

After making the line collection from Shan-si and Shen-si, 
which formed the subject of No. XI. of the present series of 
papers, the Duke of Bedford's collector, Mr. Malcolm Andei'son, 
paid a visit to Europe and America, and then I'eturned again to 
China in the autumn of 1909. He there obtained the services of 
Dr. J. A. C. Smith a,nd Mr. Kingdon Ward, who accompa,nied 
him in his fiirther explorations inland. 

The ptirty first went up into Southern Shen-si, whence a, small 
collection was sent home, which included the three species whose 
descriptions w-ere published in the 'Proceedings'*. The other 
forms then obtained will be referred to in a succeeding pnper. 

The next collection was made in the mountain I'egion of S.W. 
Shen-si, but, owing to accidental delays, havS reached London 
later than the succeeding set, which forms the subject of the 
pi-esent paper. 

Mr. Anderson's part}' then proceeded into Kan-su, a province 

* P. Z. S. 1910, p. 635. These descriptions, though not -:o nurabcved, may be 
considered to form No. XII, of the present series of papers. 



MAMMALS FiiOM WES'lERX CHINA. 159 

hitherto entii'ely uurepreseiited in our National Museum. The 
collection there made and now described is of the utmost interest 
and greatly inci'eases our knowledge of the Chinese Fauna. 

Then a move was made southward into Western 8ze-chwa.n, 
the main objective of Mr. Anderson's long journey. For it was 
in this region that the famous collections of Pere David were 
formed, and authentic specimens of the many species he dis- 
covered were vital desiderata for all satisfactory work on Chinese 
Mammals. Collections were made at and near Ta-tsien-lu, and 
again with great success on the sacred mountain Omi-san, near 
Kia-ting-fu. 

The result of the party's lal)oui\s forms, I believe, the finest 
collection of small mammals that has ever come from China. No 
less than 16 species and subspecies have had to be described as 
new*, while authentic specimens have been obtained of nearly 
all Pere David's discoveries. 

Special mention may be made of the series of Xeotetracv.s, a 
peculiar Insectivore recently described by Prof. Trouessart, of the 
black-striped Shrews, no such coloration having been previously 
described in the Family, of the specimens of Uropsilus and 
Marinella, and of the remarkable Vole Proedromj/s hedfordi, the 
type of a new genus. In all there are 360 specimens, belonging 
to 48 species and subspecies. 

The collection, as a whole, gives striking evidence of the uni- 
formity of the mammal fauna of China north of an east and west 
line at about 34° N., corresponding in its western part with the 
Tsin-ling range of mountains. Specimens from Korea on the 
east, through Shantung, Shan-si and Shen-si to Ka,n-su on the 
west, are all remarkably uniform in character, while there is 
an abrupt change on passing fi-om Kan-su to the southei-n side 
of the range into Sze-chwan. The inhabitants of the former are 
all " desert-coloured " animals, and of the latter dark Kiiimals — ■ 
no doubt because of the more forested nature of the country. 

Zoologists have every reason to be grateful to the Society's 
President for this fine addition to the available material for 
working out the Eastern Asiatic Fauna. It forms a worthy 
supplement to the results that have been already attained by the 
same means in Japan, Korea, and other pnrts of China, the whole 
constituting one of the most magnificent series of collections that 
has ever been obtained. 

Considering again the hardships that had to be undergone, the 
roughness and wild nature of the country, the unfriendliness of 
the natives, at whose hands Mr. J. W. Brooke had recently met 
his death, and the severe climatic conditions, we must extend our 
admiration to Messrs. Anderson, Smith, and Ward, for the 
extent of the collection and the excellent condition in which they 
have succeeded in sending it home. 

* The complete account of these new forms appears in this communication, but 
since the names and piTli-minavy "rlin.wnoses wpve published in the ' Abstract,' No. 90 
1911, these species are distinguished by the namrs being underlined. - -Kdjtok. 



160 MR. OI.DFIELD THOMAS OX 

1. Rhinolophus oorxutus pumilus K. And. 

c?. 2560, 2561, 2563. $. 2562, 2564. Kia-ting-fu, S. Sze- 
chwan. 1200'. 

Apparently indistinguishable from the type, with which 
Dr. Andersen has eompai'eil it. 

2. Barbastella darjelingensis Hodgs. 

c?. 2523. Omi-san. 9500'. 

A specimen of this rare species was also obtained in Sze-chwan 
— at Yin-shin-wan, N.W. of Chong-tu — by Mr. W. IST. Fergusson. 
I can find no difierence between these specimens and Hodgson's 
type from Darjiling. 

3. PlECOTUS ARIEL. 

Thos. Abstr. P. Z.S. 1911, p. 3 (Feb. 14). 

$.2343. Ta-tsien-lu, Sze-chwan. 8400'. B.M. No. 1 1.2.1.6. 
Collected 23 June, 1910. Type. 

Most nearly related to P. ivardi, but da.rker. 
■ Size among the largest of the genus. Far of medium length. 
General colour dark " broccoli brown," the ends of the hairs dull 
drab. Under surface 'paler drab, the bases of the hairs dark 
slaty. Muzzle blackish. Proportions of thumbs and hind limbs 
as in sacrimontis. 

Skull large, with swollen brain-case and broad interorbital 
reo'ion. Bulla? large, only less than those of the Egyptian species 
P. chrisiiei. 

Dimensions of the type, the starred measurements taken in the 
flesh :— 

Forearm 44 mm. 

Head and body 53* mm. ; tail 57* ; ear 43* ; third finger, 
metacarpus 39-5, first phalanx 15-5 ; thumb (c. u., exclusive of 
metacarpal) 9*2 ; lower leg and hind foot (c. u.) 31. 

Skull: greatest length 17-2; basi-sinual length 13-2; zygomatic 
breadth 9; intertemporal constriction 4 ; mastoid breadth 9-4-, 
front of canine to back of m^ 5-6 ; greatest horizontal diameter 
of bullae 4-6. 

Uab. and T'ype as above. 

Of the genus Plecotus, P. homochrous (Nepal) a,nd ptcck (Murree) 
(doubtfulfy distinct from each other) stand aside from the rest 
owing to their narrow brain-case ; auritus (Europe) has rounder 
skull, small bullpe, and short thumbs; christiei* (Egypt) large 
brain-case and very large biillfe ; vxtrdi (Ladak and Kashmir) 
large skull, rather large bullfe, long thumbs, and very pale colour ; 
sacrimontis (Japan) large skull, luther small bullre, and long 
thumbs ; and finally the present species has large rounded skull, 
large bullse, long thumbs, and dark colour. No doubt it is most 

* Althong-h the name (egi,iptiac.uis is quoted both hy Dobsou and Trouessart from 
Is, (lenffvoy's paper of 1832, I fail to find anything but " Oreilltird d'Egypte " in 
that pappi-j and therefore use Graj^'s name of 1838. 



MAMMALS FROM WESTERX CIIIXA. IGl 

nearly allied to P. loardl, but the colours of the two are iieai^ly at 
the opposite ends of the scale. 

4. Myotis ai.tarium. 

Thos. Abstr. P. Z. S. 1911, p. 3 (Feb. 14). 

(? . 2427, 2428, 2429, 2431, 2433. $ . 2422-2426, 2430, 2432. 
Omi-san, Sze-chwan. 6000'. 

A large species — about the size of M. bechsieini — -the skull 
remarkably shortened. 

Size large as compared to ordinary small species of Myotis. 
Fur rather long, but thin and poor, at least in summer pelage ; 
hairs of back about 8 mm. in length. General colour above 
uniform dull brown (paler than " Front's brown "), the tips of the 
hairs rather lighter. Under surface little lighter than upper, 
except that the tips of t]}e hairs are more distinctly lighter. Ears 
long, nearly as long as in M. bechsteini, but rather narrow ; inner 
edge evenly convex, outer slightly concave above, convex in lower 
half, a strongly develo^jed antitragal lobule at its base, separated 
by a deep notch. Tragus long, not very sharply pointed, evenly 
but slightly curved outwards ; a well-marked lobule at its outer 
base. Membranes naked throughout ; no fringe on interfemoral. 
Feet rather above normal size, but not as in " Leuconoe^' ; calcar 
extending I'ather more than halfway towards the tip of the tail ; 
a very narrow postcalcareal lobule. 

Skull different in shape from that of ordinary il/yo^is, owing to 
the shortening of the rostrum, which is broad and evenly narrowed 
forwards, instead of there being a narrow and nearly parallel-sided 
anterior portion. This condition is, however, approached in 
M. pequiniiis. Upper outline without frontal convexity, nearly 
straight from its highest point near the lambda to a point over 
the small premolars, then abruptly concave, the short nasal region 
7'eiroussee. Palate unusually vaidted. 

Middle upper premolar about half the size of the anterior, both 
in the tooth-row ; and the same is the case with the lower 
premolars. 

Dimensions of the type, the stain^ed measurements taken in the 
flesh :— 

Forearm 45 mm. 

Head and body 55* ; tail 48* ; ear 22* ; ti\agus on inner edge 8 ; 
third finger, metacarpus 40, 1st phalanx 13"3; lower leg and 
foot (c. u.) 29. 

Skull: greatest length 15-2, basi-sinual length 12; breadth of 
brain-case 7*9 ; front of canine to back of m^ 6"5. 

Mab. as above. 

Ti/pe. Adult female. B.M. ISTo. 11.2.1.9. Original number 
2423. Collected 2 August, 1910. 

This is a most peculiar species and readily recognizable by its size, 
long narrow ears, and the unusual shape of its skull, which differs 
considerably from that of most members of the genus, although 
another Chinese species, M. peqainius, shows an approach to it. 

Proc. Zool. Soc— 1911, No. XI. 11 



162 MR. OLDFIELD THOMAS ON 

5. Myotis moupinensis M.-Edw, 

S . 2322. 23 miles S.E. of Ta-tsien-lii, Sze-cliwan. 7500'. 

Milne-Edwards's Vesj^ertilio '})ipu2nnensis was placed by Dobson 
as a synonym of V. muricola Hodgs., but is I'eadily distinguished 
by the sharp notch in the outer edge of the ear. 

Two specimens of it were also obtained by Mr. W. N. Fei-gusson 
at Yin-shin-wan, IST.W, of Chong-tu. 

6. Neotetracus sinensis Trouess. 

S . 2416. 45 miles S.W. of Ya-chow, W. Sze-chwan. 
6. 2445, 2459 {jg.), 2460 (yg.), 2467, 2468, 2469, 2471, 2480. 
5 . 2434 (imm.), 2444, 2446, 2450, 2472. Omi-san. 6000'. 

This series of the highly interesting Insectivore, Neotetracus, 
recently described by Prof. Trouessai't, forms a most valuable 
addition to the Museum Collection, and at the same time enables 
me to increase our knowledge of its distribution and characteristics. 

The chief 23oint I have to note is its milk dentition, Mr. Ander- 
son's series including one half -grown and two quite j^oung 
specimens. 

The result of my examination is the following formula, showing 
both the milk and permanent teeth : — 



i 1 

• < — 

\ 1 

1 


. 2 

2 
2 

. 2 


3 (I 

1 1 

— C. < - 

3 1 1 

3 LI 


ri 
r ! 


. 




3 

3 


. 4 

4 


1 

M. — 
1 


2 

2 


3 


1 




. 


3 

3 


4 

4 


3 



Comparing this with the formulae of other members of the 
group *, we find that it is almost precisely the same as that of 
Hylomys, agreeing with it even in the absence of a third upper 
milk incisor, and the presence of the corresponding tooth below. 
The main difference is in the entii-e absence, both above and 
below, in milk and permanent series, of p'"*, this tooth being present 
in both milk and permanent series of Hylomys above, and in the 
permanent series below. 

The proportions of the milk-teeth are not very different from 
those of Hylomys, though di^ surpasses di" to a greater extent. 

While the permanent upper carnassial (p*) is of the usual shape 
characteristic of the family, its predecessor (dp^) is quite peculiar 
in the entire suppression of the small postero- internal cusp and 
the reduction of the antero-intei-nal, these being in Hylomys 
well developed and approximately similar in position to those on 
the permanent p''. They are also both present in Gymnura and 
Erinaceus. 

Taking all characters into consideration, I am of opinion that 
Neotetracus is most neai-ly related to Hylomys and is strictly a 
member of the Gymnurinpe, without any special leaning towards 
the Erinaceinee. 

* Cf. P. Z. S. 1892, p. 505. 



MAMMALS FROM WESTERN CHINA. 163 

" In damp forest ; no wnter, 
" Stomach contents, earthworms. 

" 4 embiyos — mamnife 2 — 2 = 8, another specimen with 
5 embryos."— M. P. A. 

7. Talpa longirostris M.-Edw. 
J. 2538. Omi-san. 9500'. 

An old individual from Ynng-li-pa, jST.W. Sze-chwaii, obtained 
by one of Mr. Styan's collectors, was previously the only specimen 
of this species in the Museum. 

8. Uropsilus soricipes M.-Edw. 

J. 2457, 2504, 2520, 2540, 2550, 2554. $. 2489, 2527, 
2539. Omi-san. 6000-9500'. 

Although, owing to their being occasionally deciduous, the 
teeth of this animal may have been quite correctly indicated for 
the type specimen by Prof. Milne-Edwards, the full normal 
number of teeth would appear to be greater both above and 
below than he stated. For, above, in front of the larger p^ there 
is genei'ally present a minute tooth smaller than any other in the 
jaw, and in the lower jaw there is a similarly minute tooth 
immediately following the large anterior incisor. Neither of 
these is indicated by Milne-Edwards. The full number of the 
permanent teeth would therefore seem to be 

l-l O.i, P.|, M.| = L%2=38, 

the same as in Dymecodon. In the milk-dentition, howevei^ 
there is an interesting difference, as there are three lower milk- 
premolars in addition to the non-changing p^. 

Homologising the individual teeth, the full formula would 
therefoi'e seem to be : — 

f 1 . . 3 . 4 

I 3.4 1.2.3 
I. -< C. << - P. < M. 

I 2.3.4 123 

LI . . 3 . 4 

This formula may be compared with that of Dymecodon, as given 
in a previous paper of the present series * . 

The chief difi'erences in the dentition of the two genera are the 
minute size (and occasional absence) of p^ in Uroj^sihcs, this tootli 
in Dymecodon being larger than p\ the rounder, less carnassial 
shape of p^ in Urojjsilus, and, in the lower jaw, the extremely 
minute size of the posterior of the two incisors (i.^) and a con- 
siderable superiority in size of Pg over p^, these teeth being 
subequal in Dymecodon. 

This interesting Insectivore had not been previously represented 
in the British Museum Collection. 



fl . 


2 


, 3 


f 1 


1 1 . 

1 


2 


, 3 


1 

C. << - 
1 1 


\ 

, 


. 2 , 


. 3 


^0 , 


2 


. 3 


l.^l 



* p. Z. S 1908, p. 51 (footnote). 



11* 



164 MR. OLD FIELD THOMAS OX 

9. SOREX CYLIXDRICAUDA M.-Edw. 

J . 2507, 2547. $ . 2548, 2555. Omi-san. 9500'. 

These Shrews seemed on arrival to represent a totally new form 
of the group, for they are conspicuous!)^ marked with a deep black 
line i-unning down the centre of the back, a coloration hitherto 
unrecorded in the family. 

But after a personal examination, kindly facilitated by 
Prof. Trouessart, of the species described under the above 
name by Milne-Edwards, I have come to the conclusion that 
Mr. Anderson's specimens ai'e referable to that animal, although 
its describer made no mention of the dorsal line. Probably the 
type was skinned out of spirit, and was already discoloured when 
he described it. 

The skull agrees exactly in size and dental characters with that 
of the type. 

10. (SoREX BEDFORDI^. 

Thos. Abstr. P.Z. S. 191 1, p. 3 (Feb 14). 

S . 2323, 2337. 23 miles S.E. of Ta-tsien-lu. W. Sze-chwan. 

cJ. 2359, 2374, 2391. Ta-tsien-lu. 9000'. 

c^. 2441, 2475, 2490, 2506, 2521, 2.541. $. 2496, 2497, 
2500, 2522, 2524, 2532, 2542, 2549. Omi-san. 6000'-9000'. 

A smaller species with a dorsal stripe. 

Size decidedly smaller than in S. cylindricauda, to which it 
perhaps bears the relationship that S. minutus does to S. araneus. 
Fur soft and fine; hairs of back about 4"2-4'4 mm. in length. 
General colour above brown, something between " bistre " and 
'• Front's brown"; scarcely or not lighter below. Back with an 
indistinct doisal stiipe, like that of 8. cylindricauda, but usually 
less distinct, sometimes only discernible as a faint darker median 
wash often broken over the shoulders ; in one instance, however 
(IS'o. 2532), it is as strongly black as in the larger species. Hands 
and feet pale brown. Tail dark brown above and terminally 
below, lighter proximally below. 

Skull rather smaller than that of S. araneAbs. Three anterior 
unicuspids equal; two posterior smaller and subequal, the posterior 
commonly the larger. 

Dimensions of the type : — 

Head and body 55 nun. ; tail 55 ; hind foot 13. 

Skull: condylo-basal length 17'4, greatest breadth 8"4, upper 
tooth-series 7*4 ; front of p* to back of m* 3"5. 

Hah. as above. Type from Omi-san, 9500'. 

Type. Adult male. B.M. No. 11.2.1.41. Original nimiber 
2541. Collected 19 August, 1910. 

The charactei'istic striping of this Shrew is not normally so well 
marked as in S. cylind,ricauda, but it is quite sufficiently evident 
in all the specimens to afford a ready means of distinguishing the 
species from any of its allies. 

I have named this very pretty little species in honour of the 
Duchess of Bedford, whose interest in zoology is well knoAvn. 



MAMMALS FROM NVKSTERX CHIXA. 165 

11, SOREX WARDI. 



Thos. Abstr. P. Z. S. 11)11, p. 3 (Fel). 14). 
(^.2230. Tau-cliovv, Kan-su. 9000'. 25 March, 1910. B.M. 
No. 11.2.1.46. Tyioe. 

Like *S'. bedfordice, but paler and with smaller brain-case. 

Essential characters as in *S'. bedfordice. Hairs of back 5"6 mm. 
in length. Colour paler, the general colour of the body nearly 
approaching " drab," and the tips of the hairs of the under surface 
drabby white, markedly contrasting with their slaty bases ; a line 
of demarcation pei'ceptible on sides. Dark dorsal streak well 
mai-keil, more so than in average S. bedfordice. Hands, feet, a,nd 
underside of tail brownish white, the tail much more sharply 
bicolor than in bedfordice. 

Skull with the bi-ain-case much smaller, and, especially, lower 
than in S. bedfordice, its length 4'5 mm. as compared with about 
5"5 in average specimens of that animal. Unicuspids subequal 
tliroughout, the fourth slightly the smallest. Dental pigment- 
ation moderate. 

Dimensions of the type : — 

Head and body 53 mm. ; tail 49 ; hind foot 12 ; ear 8. 

Skull: condylo-basal length 17, greatest breadth 8"2 ; upper 
tooth-series 7"4 ; front of p* to back of m^ 3"7. 

Hab. and Type as above. 

The paler colour, slightly shorter skull-length, and smaller and 
much lower brain-case will readily distinguish this Kan-su striped 
Shrew from its Sze-chwan ally. 

I have named it in honour of Mr. F. Kingdon Ward, one of 
Mr. Anderson's assistants during the expedition. 

12. SORICULUS MACRURUS Blanf. 

S. 2493. 2. 2503. Omi-san. 9500'. 
Agrees closely with Blanford's type from Darjiling. 

13. SORICULUS SACRATUS. 

Thos. Abstr. P. Z. S. 1911, p. 4 (Feb. 14). 

S. 2442, 2451, 2476. $. 2452, 2477, 2485. Omi-san. 
6000'. 

A small species allied to ;S'. ccmdatus ; tail rather shorter than 
head and body. 

Size smaller than in any described species. Form light and 
delicate. Fiu^ fine, velvety ; hairs of back about 4*5 mm. in 
length. General colour near " slate-grey," varying in some 
specimens towards brown. Under surface paler, glossy "smoke- 
grey," varying towards " broccoli-brown." Hands and feet pale 
brown. Tail a little shorter than the head and body, about as in 
S. caudatus., brown above, glossy whitish below. 

Skull markedly smaller than that of S. ccmdatus, especially in 
the size of the brain-case, the facial part, with the tooth-series, 
being nearly as long as in the allied form. Teeth as in S. cavdcctus ; 
distinguished fiom those of the last species by the tooth-i'ows 



166 Mil. OLDiaELD THOMAS ON 

being fiivther apart anteriorly, so that tlie median space between 
tlie binder lobe of the first incisors is about equal in bi'eadth to 
that lobe, markedly narrower in S. macrui-us. 

Dimensions of the type, measured in skin : — 

Head and body 60 mm. ; tail 54 ; hind foot (c. u.) 14. 

Skull: greatest length (exclusive of incisors) 18'1 ; basal 
length 15"7; greatest breadth 95 ; upper tooth-series 8"1 ; front 
of J)' to back of nr 4'4. 

Ilab. as above. 

Ty2^e. Adult § . B.M. No. 11.2.1.54. Original number 2485. 
Collected 10 August, 1910. 

This is evidently the IocmI repi'esentntive of >S'. caudatas, but is 
readily distinguishable by its much smaller brain-case. 

14. Chodsigoa smitiiii. 

Thos. Abstr. P. Z. S. 1911, p. 4 (Feb. 14). 

c?. 2349. Ta-tsien-lu. 9000'. 27 June, 1910. B.M. No. 
11.2.1.537. Ty2)e. 

Proportions as in Ch. hyiisib'm ; size much larger. 

Size veiy large, nearly equaling that of the long-tailed Vh. 
salenshii Kashtch. Fur fine and soft ; hairs of back (summer) 
3*8 mm. in length. General colour viniform "mouse-grey"; 
under surface similar in colour, though, owing to the difierent 
texture of the hairs, the surface ap})ears more glossy. Muzzle 
scarcely darker than body, Latei'al ghuid not conspicuous, nearer 
the hip than the shoulder. Hands and feet brownish white, 
lighter terminally. Tail rather shorter than head and body, pale 
brown above, whitish below. 

Skull much larger than that of Ch. hi/psihia, the brain- case 
flattened, heavily ridged. Proportions of teeth as in that species, 
their tips as usual lightly pigmented. 

Dimensions of the type, an old male : — ■ 

Head and body 85 nun. ; tail 68 ; hind foot 18. 

Skull: condylo-basal length 22-5, greatest breadth 10-5; ujjper 
tooth-series 10*2 ; front of p' to back of nr 5"3. 

JIah. and Type as above. 

This fine species is conspicuously lai'ger than its ally Ch. h>/p- 
s/6i» de Wint., with Avhich I synonymize CJi. herezovshi Kashtch. 
It is rather smaller than the long-tailed Ch. salenskii Kashtch. 

I have named it in honour of Dr. J. A. C. Smith, who ac- 
companied Mr. Andei'son during the Sze-chwan trip, and assisted 
largely in making the collection. 

15. Blarinella quadraticauda M.-Edw. 

J . 2505, 2556, 2558. $.2557. Omi-san. 9500'. 

Blarinella, g. n. 

More allied to the N.-American Blariva than to any of the 
Old World genera of Shrews. Ears very small, though neither 



MAMMALS FROM WESTERN CHINA. 167 

SO ruflimentaiy nor so hair-covered as is tlie case in Marina ; 
their conch wholly below the surface of the fur. Tail shoi't, but 
not nearly so shoi't as in Blarina. Claws large, the anterior 
inconspicuously larger than the posterior. 

Skull in genei'al build about as in the more delicate species of 
tlic sul)genus Cryptotis^ though the rostrum is hardly so high. 

Teeth numbering 32, as in Sorex and true Blarina. Upper 
unicuspids normally 5, but of such proportions and so slanted that 
only 3 are as a rule visible in a side view, as in Cryiitoiis. First 
two large and subequal, third about half their size, its hinder 
edge level with the front of p'; fourth flattened from before 
backwards, about half the size of third, and jammed close against 
it, between it and the minute fifth, which is entirely in the inner 
half of the tooth-i'ow, close against the large p'. Molars S(iuare, 
practically without intei'spaces posteriorly. Lower teeth about 
as in Blarina, the large incisors with thi-ee notches. All teeth 
heavily pigmented, as in Blarina. 

Type, B. quadraticauda {Sorex quadraticauda M.-Edw.). 

The discovery that the Blarina group is represented in the Old 
World is a most interesting one and very similar to that of the 
Sze-chwan Zapus, published by M. Pousargxies in 1896. The 
present species was described as a Sorex by Milne-Edwards, but 
when examining the type in Pax-is some years ago I felt sure 
it was either Blarina or neai'ly allied to it, and Mr. Anderson's 
series now enables me to place it with certainty. 

Of the four specimens obtained thi'ee have five unicuspids, as 
above described, but in one (No. 2505) the minute fifth is absent 
on both sides, so that it seemed possible the type was in the same 
condition, only four unicuspids having been described and figured. 
A i'e-examina.tion of it, which I owe to the kindness of Prof. 
Trouessart, shows that it presents the intermediate condition of 
having on the right side only 4 unicuspids, the 5th minute one 
being missing, just as in no. 2505, while on the left this tooth 
is present. Owing, however, to the accidental loss of the 3rd and 
4th unicuspids on this left side, the fact that the minute tooth 
still present, lying pressed against p'', corresponded to the fifth 
tooth that we now know to be normal to the species, had not 
been previously observed. There is therefore no doubt whatever 
as to the specific identity of the pi-esent series with Milne- 
Edwards's animal. 

The four groups of short-tailed Shrews may be i-eadily dis- 
tinguished from each other by the number and relative visibility 
from outside of the upper unicuspids, as follows : — 

5 unicuspids, 4 visible from outside. Blarina. 

5 ,, 3 ,, ,, Blnrinella. 

4 „ 3 „ ,, Cryptotis. 

3 ,, 3 ,, ,, Notiosorex. 

So entirely are both posterior unicuspids generally hidden in 
Blarinella that the figure of the teeth of B. {Cryptotis) parva 



168 MK. OLDFIELD THOMAS ON 

given by Merriam in his Monograph* would almost do for it, 
were it not for the much greater size and posterior extension of 
the large upper incisor of the Asiatic animal. 

16. Crocidura attenuata M.-Edw. 

d . 2279. 2 • 2278. Wen-hsien Country, Kan-su. 

17. Anourosoeex squamipes M.-Edw. 

J . 2407, 2410, 2412. $ . 2408, 2409, 2411. 45 miles W. of 
Ya-chow, Sze-cliwan. 

d. 2415, 2417, 2420. $. 2418, 2412. 45 miles S.W. of 
Ya-chow. 

c?. 2435, 2436, 2458, 2464, 2470, 2481, 2485, 2487, 2501, 
2533. 2 • 2440, 2455, 2465, 2483, 2486, 2488, 2502. Omi-san. 
6000-9500'. 

The first-mentioned place would appear to be about 50 miles 
S.W. of Moupin, the type locality of the species. 

18. YULPES sp. 

2 . 2201. Near Si-Ho-Hsien, S. Kan-su. 

Apparently representing V.filchneri Matsch. The relationship 
of this form to the Moupin Fox (F. hooly David) is not clear. 

19. MUSTELA RUSSELLIANA. 

Thos. Abstr. P. Z. S. 1911, p. 4 (Feb. 14). 

c?. 2381,2382. 2.2388,2389,2390. Ta-tsien-lu. 12,000'. 

A very small species, with the contrasted body-colours and 
tuftless tail of M. kathiah. 

Size extremely small, about that of the veiy smallest known 
Weasel — the American M. rixosa Bangs. Colours of upper and 
lower surfaces sharply contrasted, as in M. katliiah and the Stoats, 
not as in M. nivalis. Upper surface uniform dark bi'own, rather 
less rich and more drabby than in M. kathiah. No darker 
markings on face or ears. Under surface a beautiful pinkish 
buff, turning into white anteriorly on the chin, interramia, and 
lips. Line of demarcation veiy sharply marked, running from 
upper lip to ankle. Arms brown externally, bufiy on inner 
aspect ; hands brown, sometimes mottled with white ; hind limbs 
similarly brown externally and bufFy on the inner aspect ; palms 
and soles densely hairy. Tail piopoi'tionally shorter than in 
AI. kathiah, slender, not tufted, uniformly bi^own, the tip not 
noticeably darker. 

Skull distinguishable from that of all other allied forms by its 
extremely small size. 

Dimensions of the type : — 

Head and body 133 mm. ; tail 54 ; hind foot 22 ; ear 11. 

Skull : condylo-basal length 29-3 ; basal length 27"2 ; zygomatic 
breadth 15-2 ; interorbital breadth 6-2 ; breadth of biain-case 14 ; 

* N. Am, Fauiirt. no. 10. pi. iii. fip'. 2, 1805. 



MAMMALS FUOM WESTERN ClilXA. 169 

pn.lfita.l length 11 ; front of canine to back of molar 8 ; ji'' on outer 
edge 2-9. 

A male specimen measure-; : liead and body 138 mm. ; tail 54 ; 
bind foot 24; carnassial tootb 3-1. 

Hah. as above. 

Type. Adult female. B.M. No. 11.2.1.86. Original number 
2388. Collected 1 July, 1910. 

This most beautiful little Weasel competes with the American 
M. rixosa Bangs for the honour of being the smallest existing 
Ca.rnivore, and is readily distinguishable from any Old World 
Mitstela by its small size alone. Its essential coloration is very 
much as in the much larger M. kathiah, of which a local luce 
(dorsalis) has been recorded from this region. But the hind foot 
of that animal measures 30 mm., so that it is evidently quite a 
distinct species. 

The combination Mastela hedfordi being already in use, I have 
given to this striking new species a designation ba.sed on the 
family name of the donor of the present magnificent accession to 
the National Collection of Mammalia. 

20. Tamiops swinhoei M.-Edw. i 
d'. 2510. Omi-san, Sze-chwan. 9500'. 

21. Sciurotamias davidianus M.-Edw, 

5 . 2277. 40 miles N. of Kai-chow, S. Kan-su. 

22. Sciurotamias davidianus ooxsobrixus M.-Edw. 

J . 2320, 2321. $ . 2316. 23 miles S.E. of Ta-tsien-lu. 7500'. 
6. 2456. Omi-san. 6000'. 

The Ta-tsien-lu specimens are not very typical consobrinus, but 
are none of them adult. 

23. Marmota robusta M.-Edw. 

5 . 2270. Old Tau-chow, Kan-su. 9000'. 
?. 2269. Near Tau-chow. 10,000'. 

24. Mus GANSUEXsis Sat. 

6 . 2220, 2221, 2222. 10 milts S. of Tau-chow, Kan-su. 
Described as a Leggada by Satunin. 

25. Epimys coxfuciaxus luticolor Thos. 

c^. 2196, 2197, 2199,2200. §. 2198. Near Si-Ho-Hsien, 
S. Kan-su. 5000'. 

c? . 2206, 2219. 60 miles S.E. of Min-chow, Kan-su. 8000'. 

6. 2280, 2286, 2287, 2294, 2295, 2301, 2302. 2309, 2311. 
$. 2281, 2282, 2288, 2310, 2312. Wen-hsien ' Country, S. 

Kan-su. 

These Kan-su specimens tend to be, as is right, intermediate 



170 MR. OLDFIELD THOMAS ON 

between the true luticolor of Northern Shen-si and the darker 
covfacianus of Sze-chwan. The hghter specimens among tliem 
absohitely match some of the typical seiies, while the darker ones 
match the lighter individuals of the next following set. 

2G. Epimys confucianus M.-Edw. 

d. 2313, 2318. $.2314,2315,2317,2319, 23 miles S.E. of 
Ta-tsien-lu, Sze-chwan. 

J . 2370, 2371, 2372. Ta-tsien-lu. 9000'. 

S . 2405. 2 • 2406. 18 miles E. of Ta-tsien-lu. 5500'. 

5 . 2413. 45 miles W. of Ya-chow. 

(5 . 2443, 2453, 2462, 2463. Omi-san. 6000'. 

Pending the arrival of tojjotypes from Ohong-tu, these speci- 
mens may be accepted as typical of the true confucianus. 

27. Epimi's ling Bonh. 

9. 2283,2289. Wen-hsien Country, Kan- su. 

These beautiful ochraceous-buff specimens are quite like the 
typical series from Kuatun, and are also very similar to an 
example in the Museum from Sikkim, referred — and perhaps 
correctly — to E.jerdoni Blyth. 

But as this latter determination needs revision with better 
material, I use the Chinese name which happens to be 
available. 

28. Epimys excelsior. 

Thos. Abstr. P.Z. S. 1911, p. 4 (Feb. 14.) 
cJ. 2360, 2379. $.2380. Ta-tsien-lu, Sze-chwan. 9000'. 
S. 2511, 2526, 2545. Omi-san. 9500'. 

General characters of E. confucianus, but size larger. 

Size larger than in confacianus, proportionate length of tail 
about the same. Fur long and rather coarse, not mixed with 
spines in any of these specimens ; hairs of back about 14 mm. in 
length, the isolated bristle-hairs attaining 20. General colour of 
adults coarsely lined cinnamon or clay-colour — not unlike bright- 
coloured specimens of E. norvegicus. Sides clearer and more buffy. 
Belly pure sharply defined white or creamy wdiite, which extends 
anteriorly well up over the upper lip. Top of muzzle greyer. 
Eyes with darker rims. Ears of medium length, greyish brown. 
Hands white above. Feet white with prominent dark metapodial 
patches. Tail rather longer than head and body, well-haired and 
tufted, brown above for about two-thirds its length, white at the 
end and below ; hairs at end 7-8 mm. in length. 

Skull differing from that of confucianus by its larger size and 
more delicate build. Nasals long and slender. Interoi'bital 
region narrow, its borders sharp-edged, but without definite 
beads. Brain-case smooth, markedly convex above. Anteorbital 
pla.te but little projected. Palatal foramen long. Bulte small. 
Molars decidedly larger than in confaciamcs. 



MAMMALS FROM WESTERN CHINA. 171 

Dimensions of tlie type : — 

Head and body 178 nun. ; tail 193 ; hind foot 30 ; enr 23. 

Skull : greatest length 39 ; basilar length 30"3 ; zygomatic 
breadth 18; nasals 15-2 x 4"3 ; interorbital breadth 4-3 ; breadth of 
brain-case 15*6; palatilar length 17; palatal foramina 7"5 ; upper 
molar series 6'8. 

Ilah. of type. Ta-tsien-lu. 9000'. 

Type. Adult male, B.M. No. 1 1.2.1.131. Original number 2379. 
Collected 30 June, 1910. 

The true Rats of the genus Epimys are, as usual, some of the 
most difficult of the collection to make out satisfactorily, owing 
to their variability in both colour and skull-charactei-s. But it 
seems clear that this foi-ni from the high ground of Ta-tsien-lu 
and the upper part of Omi-san is a distinct species from the 
widely spread E. confacianus, differing by its greater size, longei- 
skull, and, in fully adult specimens, its paler coloui- — at least, as 
compared with the typical dark E. covfucianus of W. Sze-chwan. 
It is, of course, a member of the saine group of the genus, ns is 
also the next species, which is again a step larger in size than 
E. excelsior. 

29. Epimys andersoni. 

Thos. Abstr. P. Z. S. 1911, p. 4 (Feb. 14). 

$.2478. Omi-san. 6000'. Collected 8 August, 1910. B.M. 
No. 11.2.1.135. Type. 

Like E. confucianus and excelsior., but very much larger and 
with longer tail. 

Size large, though not equaling the giant of this group, 
E. edwardsi. Fur coarse, not spinous ; hairs of back about 10 mm. 
in length. Greneral colour as in E. excelsior, a dull-lined clny- 
colour, more bufty on the sides. Under surface pure sharply 
defined white. Top of muzzle greyish ; sides dark brown, this 
colour passing backwards and joining the dark eye-rings. Ears 
of medium length, the proectote dull blackish. Hands white, the 
dark of the forearms encroaching a little on the metacarpus. 
Feet brown over the greater part of the metatai^sus ; toes and 
hallucal border of metatarsus white. Tail very long, faii-ly well- 
haired, not conspicuously tufted, brown for its proximal half 
above, then changing gradually to white ; underside wholly white, 
except just at the base. 

Skull in general structure like that of E. excelsior, but larger 
throughout ; brain-case rather less convex above. 

Dimensions of the type (young ;idult) : — 

Head and body 164 mm. ; tai'f 248 ; hind foot 37 ; ear 26-r,. 

Skull : greatest length 43 ; basilar length 35; nasals 16'3 ; breadth 
of brain-case 17-2 ; palatilar length 19-8; palatal foramina 8-5; 
upper molar series 8 '2. 

The dimensions will at once separate from any of its allies this 
fine long-tailed Rat, which I have much pleasure in dedicating to 
its discoverer, Mr. Malcolm Anderson. 



172 MK. OLBFIELD THOMAS OX 

30. Apodemus speciosus pekinsul.e Thos. 

c^. 2202, 2211, 2212, 2213, 2218. 60 miles S.E. of Min-diow, 
Kan-su. 8000', 

J. 2227,2228, 2231, 2232. $. 2229. 10 miles S. of Tau- 
chow, Kan-su. 8000'-9000'. 

d. 2234, 2235, 2237. ? . 2242. Mts. S.E. of Tau-chow, 
Kan-sn. 10,000'. 

J . 2285, 2305, 2308. Wen-hsien Country, S. Kan-su. 

1 have already commented on the resembla,nce of the Shan-si 
Field-Mouse to that of Korea, and now find that those from Kan- 
su are again similar in every respect and should undoubtedly be 
referred to the same subspecies, the animal therefore presenting 
a noteworthy example of the uniformity of the Fauna of China 
N. of lat. 34°. As usual, the representative southern form in 
Sze-chwan is at once distinguishable by its much darker colour. 

31. Apodemus speciosus chevrieri M.-Edw. 

2 c?, 1 2- 23 miles S.E. of Ta-tsien-lu, Sze-chwan. 10,000'. 
13 c?, 13$. Ta-tsien-lu. 9000'. 

17 J, 11$. Omi-san. 6000'-9500'. 

32. AroDEMUs agrarius pallidior Thos. 

(^.2195. Si-Ho-Hsien, S. Kan-su. 5000'. 

cJ . 2292. $ . 2293. Wen-hsien Country, S. Kan-su. 

33. Apodemus fergussoni. 

Thos. Abstr. P.Z. S. 1911, p. 4 (Feb. 14). 

$ . 2296. Wen-hsien Country, S. Kan-su. 

$ . 2499 (yg.), 2508, 2509 (yg.). Omi-san. 9500'. 

Near A. agrarius, but without the slightest trace of a dorsal 
line. 

Size and proportions about as in agrarius. Fur short and fine, 
without spines in a May specimen, with them in August 
examples ; hairs of back about 7 mm. in length. General colour 
a,bove uniform drab-brown, not buffy on sides. Under surface 
soiled greyish, the bases of the hairs slaty, the tips greyish white ; 
lateral line of demarcation fairly well defined. Back without the 
slightest trace of a median dorsal line, or even of a median darker 
dorsal area, some trace of a line being always perceptible in 
A. agrarius. Ears short, coloured like the head. Hands and feet 
white. Tail rather shorter than head and body, brown above, 
dull whitish below. 

Skull about as in A . agrarius. 

Dimensions of the type : — 

Head and body 107 mm. ; tail 98 ; hind foot 24 ; ear 15. 

Skull: greatest length 27-7; breadth of brain-case 12 ; upper 
mola,r series 4. 

JIah. S. Kan-su and Western Sze-chwan. 



MAMMALS FROM WESTERX CHIXA. 173 

Type. Adult female, B.M. No. 11.2.1.180. Original number 
2296. Collected 8 May, 1910. 

This Mouse is readily distinguishable from any of the forms of 
agrariiis by the entire absence of a dorsal line, some trace of a line 
being perceptible in all the available specimens of A. agrarius 
ningpoensis, the form in which it is least developed. Even then 
I should like to call it a subspecies of agrarius, but do not 
venture to do so without seeing intermediate specimens. The 
type was obtained at or near the same locality as a well-marked 
example of A. agrarius pallidior (no. 2292), so that in this region 
at least there seems to be iio intergi'adation, 

I have named this animal after the Rev. W. N. Fergusson, 
from whom the Museum has received four specimens of it, 
collected at Yin-shin-wan, N.W. of Chong-tu, in 1909. 

34. Cricetulus andersoxi Thos. 

d. 2215, 2216. $.2217. 60miles S.E. of Min-chosv, Kan-su. 
2 . 2225, 2233. 10 miles S. of Tau-chow, Kan-su. 

35. MiCEOTUS IRENE. 



Thos. Abstr. P. Z. S. 1911, p. 5 (Feb. 14). 

S. 2383, 2386, 2397, 2400, 2404. $. 2384, 2385, 2392, 
2395, 2396, 2398, 2399, 2401, 2402, 2403. Ta-tsien-lu, W. 
Sze-chwan. 9000-12,000'. 

Rather larger than M. arvalis. M^ with the same spaces con- 
nected as in Fitymys. 

Fur soft, fine, rather woolly ; hairs of back about 9 mm. in 
length. General colour above brown, under surface slaty 
washed with greyish white ; not sharply defined lateiuUy. Ears 
projecting beyond the fur, uniformly brown. A small lateral 
gland present on each side in front of the hips in the males. 
Hands and feet brownish white ; soles with six pads. Tail about 
twice the length of the foot ; brown above, paler on sides and 
below, not so strongly contrasted as in M. malcohni. Mammae 
2—2 = 8. 

Skull of the usual flattened shape, its upper outline straight or 
even slightly concave over the orbits. Brain-case remarkably 
small in proportion to the face, smooth, rounded, its ridges and 
angles scarcely developed ; muzzle slender, scarcely bowed down- 
wards ; interorbital space unusually narrow, smoothly rounded. 
Palatal foramina fairly long. Posterior palate normal, the lateral 
pits not specially deep ; mesopterygoid fossa of normal breadth, its 
anterior border rounded. Bulhe decidedly smaller than in most 
species of this size. 

Incisors of medium strength, considerably bevelled in front. M^ 
with pattei-n very similar to that figured by Blasius in M. savii 
(Faun. Deuts. fig. 221, p. 394). M^ with only four closed spaces, 
the fifth and sixth (counting from behind and including the 
posterior transverse space) opening into each other (as in Fitymys) 



174 MR. OLDFIELD THOMAS ON 

and also into the seventh, the anterior space, which is curved 
inwards to form a fifth internal angle, while there are only three 
external angles ; the tooth is thus very like that figured by 
Biichner in M. limnophilus (Mamm. Przewalsk. pi. xviii. fig. 13). 

Dimensions of the type : — 

Head and body 98 mm. ; tail 37 ; hind foot 18 ; ear 13. 

Skull : condylo-basal length 24-2 ; basilar length 22 ; greatest 
breadth 14-8 ; nasals 7 ; interorbital breadth 3'8 ; breadth of 
brain-case 12 ; height of forehead from alveolus of m" 7 ; palatilar 
length 13; palatal foramina 4-8 ; diagonal diameter of bulla in 
horizontal plane 6'5 ; upper molar series (crowns) 5-7. 

Hah. as above. 

Type. Old female, B.M. No. 11.2.1.195. Original number 2396. 
Collected 2 July, 1910. 

This species is peculiar for the proportionately small size of its 
brain-case and its narrow interorbital space. From M. limnophilus 
Biichn., which may possibly be allied to it, it is distinguishable by 
having only three internal angles on m^ ; the shape of the brain- 
case would also seem to be more normal in that animal. 

36. MiCRO TUS MALCOLMI. 

Thos. Abstr. P. Z. S. 1911, p. 5 (Feb. 14). 

S. 2245, 2250, 2251, 2252. $. 2246, 2253, 2254, 2255, 
2256, 2257. Mts. S.E. of Tau-chow, Kan-su. 

A medium-sized Yole with high arched skull, somewhat similar 
on a smaller scale to that of M. ccdamorum Thos. 

Fur soft and fine, of medium length ; hairs of back about 
9-10 mm. in length *, therefore conspicuously shorter than in 
Proedromys. Colour above lined brown, rather lighter than 
" bistre " ; under surface washed with greyish white, rather 
mai-kedly contrasted with the upper colour. Ears slightly 
projecting from the fur, brown. Upper surface of hands and feet 
dull whitish ; soles with six pads. Tail rather more than double 
the length of the foot, brown above, dull white on sides and 
below. Mammee apjjarently 2 — 2 = 8, but this cannot be made out 
with certainty. 

Skull rather like that of M. ccdamorum on a small scale, 
though not so strong and heavily ridged. In general shape it is 
not tiattened above, but high, arched above, the nasal profile 
bowed downwards, the waist narrow and proportionately far 
forward. Surface everywhere smooth and unridged ; squamosal 
projections scarcely perceptible. Nasals nearly attaining to the 
level of the premaxillse behind. Interorbital space narrow, 
smooth, evenly convex above, not flattened or hollowed mesially. 
Brain-case smoothly rounded. Palatal foramina of equal breadth 
throughout, their length exceeding that of m^^-m^ Posterior 
palate with very deep lateral pits and unusually narrow meso- 
pterygoid space. Bullae decidedly larger than the normal. 

* In tliis measurement I alwaj's ignore tlie outstanding tips of the longer liairs, 
and only measure to the ends of the thick mass of under-fur. 



MAMMALS FROM WESTERN CIIIXA, 175 

Upper incisors narrow, considerably lievelled latei-nlly. 

Molars of normal structure, the pafcterii of tlie upper ones, so 
far as spaces and angles are concerned, essentially as figured by 
Blasius (p. 379) in M. arvalis, though the posterior lobe of m'' is 
rather narrower and more elongated. Lower teeth as figured by 
the same author (p. 366) in M. rattiaqjs, m^ with a very similar 
anterior lobe and the same number of closed triangles. 

Dimensions of the type : — 

Head and body 103 mm. ; tail 40 ; hind foot (c. u.) 17 ; ear 13, 

Skull : condylo-basal length 26*2 ; basilar length 23*5 ; greatest 
breadth 15 ; nasals 7 ; interorbital breadth 3"7 ; breadth of brain- 
case 12 ; height of crown from alveolus of m'"^ 9'2 ; palatilar length 
13'2 ; diastema 8'3 ; palatal foramina 5 ; diagonal length of bulla 
in horizontal plane 8"2 ; upper molar series (crowns) 6'1. 

Other specimens are smaller, the smallest adult having a 
condylo-basal length of 23'4, with a hind foot of 16 mm. 

Hab. as above. 

Type. Adult male, B.M. No. 11.2.1.199. Original number 2245. 
Collected 4 April, 1910. 

I can find no described species to which this Vole can be 
assigned. Its high skull separates it at once from the great 
mass of flat-skulled Yoles, the shape more recalling that in 
Af. calamontm, a species possessing a lateral gland and only five 
foot-pads. Satunin's three species from Zaidam, W. of Kan-su, axe 
all markedly larger than M. mcdcolmi. Buchner's M. limnophUns 
from the same district is perhaps more closely allied, but would 
seem to have a paler general colour and a much more roughened 
and angular skull. 

37. MiCROTUs (Anteliomts) chinensis Thos. 

c?. 2328, 2329, 2332, 2338, 2339. $. 2326, 2327, 2331, 
2333, 2334,' 2335, 2340, 2341. 23 miles S.E. of Ta-tsien-lu. 
10,000'. 

cf. 2512, 2519, 2534, 2543, 2544. 9. 2491, 2492, 2528, 
2551, 2553. Omi-san. 9500'. 

This striking Vole was described in 1891 * from a specimen 
found in the stomach of a snake. No other examples have been 
obtained until now. 

The type-locality was Kia-ting-fu, quite close to Omi-san. 

38. MiCROTUS (CaRYGMYS) EVA. 

Thos. Abstr. P. Z. S. 1911, p. 4 (Feb. 14). 
d. 2238, 2239. o. 2240, 2241. Mts. S.E. of Tau-chow, 
Kan-su. 10,000'. 

[Caryomys, 

Like Eothenomys in external and general ci^anial characters, 
but the teeth with the triangles nearly all closed, instead of being 
mostly open and connected with each other. Additional postero- 
* Ann. Mng. N. H. (6) viii p. 117, 1891. 



176 MR. OLDFIELD THOMAS ox 

internal lobes ou m^ and m'^ veduced to minute and scarcely 
perceptible projections. 

Type, Microtios Inez Tlios. , 

Now that a third species, essentially similar to M. inez and 
M. nux, has turned up, I think it advisable to form a special 
subgenus for the reception of the three. On first describing 
M. inez * I purposely delayed making a new subgeneric name 
until we had evidence as to the existence of other species allied 
to that peculiar species, but full details were given as to its 
characteristics.] 

Allied to M. {C.) inez and nux, but with much longer tail. 

Form slender, the general proportions, chiefly owing to the 
unusually long tail, looking very different from those of the allied 
species. Fur long, soft, and fine ; hairs of back neai-ly 10 mm. in 
length. General colour above pale brown, greyer than in nux 
and inez, the type nearly approaching " broccoli- brown," but other 
specimens of a warmer tint. Under surface rather variable, 
greyish slaty, washed Avith buffy whitish or drab. Ears about as 
long as the fur, and of about the same colour. Hands and feet 
white ; soles with six pads. Tail remarkably long for a Vole, 
nearly as long as the body without the head ; dark brown above, 
dull whitish below. 

Skull very like that of M. nux, except that it is smoother and 
more lightly built ; but even this may be a question of age. 

Teeth almost exactly as in M. nux ; second and third spaces of 
m^, and first and second of m^, communicating with each other — 
all the other triangles closed (except, of course, in the case of m^). 

Dimensions of tlie type : — 

Head and body 88 mm. ; tail 50 ; hind foot 16-5 ; ear 12. 

Skull : condylo-basal length 22'6 ; basilar length 20'5 ; gi-eatest 
breadth 13'5; nasals 6'8 ; interorbital breadth 4; breadth of 
brain-case 11*2 ; palatal length 10*6 ; palatal foramina 41 ; upper 
molar series (crowns) 5. 

Hah. as above. 

Type. Adult male. B.M. No. 11.2.1.223. Original number 2238. 
Collected 3 April, 1910. 

This Yole is at once distinguishable from all its allies by its 
remai'kably long tail. 

39. MiCROTUS (EOTHENOMYS) MELANOGASTER M.-Edw. 

S. 2284, 2290, 2291, 2297, 2304, 2306, 2307, $. 2298, 
2299, 2300, 2303. Wen-hsien Country, S. Kan-su. 

c5' . 2421 (melanoid). 45 miles S.W. of Ya-chow, Sze-chwan, 
4000'. 

$. 2461. Omi-san. 6000'. 

I fail to find any tangible difference between thti Kan-su series 
and the two specimens from Sze-chwan, which la'tter are nearly 
topotypical of the species. 

* P. Z. S. 1908, p. 970. 



mammals fkom western ciiixa. 177 

40. Proedromys bedfordi. 



Thos. Abstr. P. Z. S. 1911, p. 4 (Feb. 14). 
2- 2214. 60 miles S.E. of Min-Chow, Kan-su. 8000'. 
11 March, 1910. B.M. Ko. 11.2.1.235. Type. 

Proedromys *. 



General external form as in Microtus. Far very long. Sole- 
parls 6. Mamm® 2—2=8. 

Skull heavily built, high, its upper profile strongly curved. 
Postorbital squamosal pi'ojections strongly developed, almost 
peg-like. Palatal foramina long. Posterior palate normal. 
Bullpe rather large. 

Teeth. Upper incisors heavy, strongly curved, not thrown 
forward ; their front face grooved on its outer third. Lower 
incisors short posteriorly, ending at the bottom of the notch 
between the angular and condyloid processes, as in Anteliomys. 
Molars with their enamel spaces all completely se|3arated from 
each other, the anterior wall of each upper space and posterior of 
each lower one unusually strongly curved, so as to form in many 
cases nearly half of a circle. Number of spaces and salient angles 
as follows : — ■ 

-ii/r -I J 5 spaces, 3 inner and 3 outer angles. t«,t 9 J 4 . 2 . 3 _ -n /r o 



5.3.3 [3.3.2 

M^ very peculiar, consisting of the usual transverse, second and 
third spaces, and then a circular posterior lobe directed externally, 
the posterior lobe of this tooth in every other Vole, so far as I am 
aware, being dii-ected inwards. Anterior space of m, forming 
a simple crescent. Mg with broad connected spaces across it as 
usual. 

I fail to find any group into which this long-haired Vole can be 
fitted. Its heavy bowed skull, grooved incisors, jseculiar curved 
walled tooth-spaces, and curious nig seem together to distinguish 
it from any described genus. The grooving of the incisors is 
possibly an individual chai-acter, but the other features of the 
animal are so marked that it should certainly have a sjiecial 
name. 

Proedromys bedfordi. 

Size medium. General form that of an average Vole, the tail 
rather more than a third the length of the head and body. Fur 
excessively long, soft, and fine; hairs of back about 16 mm. in 
length. General colour above coai'sely lined dull brown (darker 
than " broccoli- orown "), the lower flanks more drabby ; under 
surface slaty drrb washed with brownish white. Ears hairy, pale 

* Trpoei'pos, a Piosident. Named in honour of the Society';- President, to whose 
generositj' the discovery of this animal is due. 

Proc. Zool. : oc— 1911, No. XII, 12 



1,7B »TR.. OLDFIELD THOMAS OX 

brown. H:anrl& and feet dull white. Tail well-haired, brown 
above, dull white on sides and below. 

Skull and teeth as indicated above. 

Dimensions- of the type : — 

Head and body 103 mm. ; tail 41 ; hind foot 18 ; ear 13. 

Skull : basal length (c.) 26 ; greatest breadth 16 ; nasals 
7r-6x3-2; interorbitai breadth 3-6; breadth of brain-case 12-2; 
palatilar length 13-7; diastema 8; palatal foramina 6; upper 
molar series (ci^owns) 6'8.. 

Hab. and Type as above. 

Interesting and pecvilian as this species is in essential characters, 
it is externally a very ordinary-looking Yole, and indeed scarcely 
distinguishable from Microtus vialcolmi, which Mr. Anderson found 
ill some numbers in the s-aine region,. 

41.. MYOSPAiiAX cANSus Lyon. 

d. 2263, 2272, 2274. ?■.. 2262, 2264, 2265, 2266, 2267, 
2268, 2271, 2275, 2276.. 10 miles S. of Tau-chow, Kan-su. 8500'. 

These are practically topotypes- of Dr. Lyon's species, and a 
study of them shows that the series obtained in Shen-si by 
Mr.. Anderson, and previously referred to M. ccmsus'*, should 
be subspecifically separated. 

The form may be called : — 

MyOSPALAX CANSUS SHENSEIUS. 

Thos..Abstr. P..Z. S. 1911, p.. 5 (Feb. 14). 

General characters as in true ccmsus, but the colour wai'mer and 
the tooth-row longer.. 

Taking only full-coloured specimens into comsideration the 
coloured ends of the hairs above approach " ochraceous-butf," 
while in cansus they average distinctly paler, approximating to 
" pinkish-bufF." As to the tooth-row, among seven adult specimens 
of true 6■a«s^(s (including three well-grown males)the alveolar length 
of the upper series never exceeds lO'O mm., while of 16 specimens 
of she7iseius some (males) may almost reach 12-0 mm., while nearly 
all, including females, are ITO and upwards. One fully adult 
specimen only, a female, has the dimension 10"0. 

Dr. Lyon's type, a male (although originally called a female), 
seems to have been unusually large, as its tooth-row mea,sures 
about that of a female sheiiseius, and therefore more than any 
of our specimens of the Kan-su form. This Avould, however, 
appear to have been an exceptional specimen. 

Dimensions of the type : — 

Head and body 178 mm. ; tail 54 ; hind foot 30. 

Skull: condylo-basal length 45 ; zygomatic breadth 34; nasals 
18x7"7; interorbital breadth 6'5 ; palatilar length 23*5; upper 
molar series (crowns) 11-4, (alveoli) 11-9. 

Hah. Shen-si. Type from Yu-lin-fu. 

* P. Z. S. 1908, p. 978. 



MAMMALS FROM WESTKEX CHIXA. 179 

T}ipe. Old male. B.M. Xo. 9.1.1.216. Original number 1800. 
Collected 30 April, 1908. 

Dr, Allen's " Mijotalpa ''" rufescens * woiild seem also to have 
the small teeth of ti-ue M. cansus, and is* geographically closer t(i 
the latter. 

42. Rhizomys vestitus M.-Edw. 

$. 2559. Omi-s:m. 9500'. 

The relationship of this form to Gray's li. sinensis has not 
hitherto been satisfactorily defined. It seems, however, to be 
readily characterized by the much larger size of the skull, the 
great development of the cranial crests, the length and softness of 
the fur, and the shortness of the tail. li. sinensis probably came 
from somewhere in South China, perhaps the neighbourhood of 
Canton. 

K tliird Chinese species may be described as follows:- — 
Hhizomys davidi. 

Thos. Abstr. P. Z. S. 1911, p.. 5 (Feb. 14).- 

Colour and character of fur, and length of tail, about as in 
R. vestif/us ; but size much smaller, neaidy as in JR.- sinensis. 

Skull slightly larger than that of R.-sinensis, the male R. sinensis 
about equaling the female R. davidi. Crests I'ather more 
strongly developed, comparing like sex with like. Xasals more 
nariowed behind and premaxillary processes proportionately 
broader, so that, while in R: sinensis the combined breadth behind 
of the two nasals about equals either fronto-premaxiilary suture, 
in R. davidi the length of each suture is about twice the distance 
that separates their nearest points in the middle line.- Bullae 
largei'. 

Dimensions of the type, measured in skin :■ — 

Head and body (c.) 350 mm. ; tail 69 ; hind foot (wet) 42 ; ear' 
(wet) 15. ' 

Skull : condylo-basal length 66 ; upper length fi^om lambda to 
tip of nasals 53-5 ; greatest breadth 49*5 ; nasals 23*2 x 8'4 ; 
fronto-premaxiilary suture 6'5 ; interoi'bital breadth 9*2 ; greatest 
occipital breadth 30; palatilar length 35 ; diastema 20 ; upper 
molar series (crowns) 13'3. 

An old male skull measures 49 mm. in upper length. 

JTab. Kuatun, N.W. Fokien. 3500'. 

Ti/pe. Old female. B.M. No. 96.12.1.6. Original number 9. 
Collected December 1898, and presented by J. de La Touche, Esq.- 

Eleven specimens exaniined. 

This fine species, of which the Museum possesses a good series,- 
presented by Messrs. F. W, Styan, J. de La Touche, and C. B. 
Rickett, is distinguishable from R. sinensis by its soft fur, shoiter 
tail, and the above-mentioned cranial characters. From R. restitag, 
again, by its very much smaller size. 

* Bull. Am. Mus. N. 11, x.vvi. p. 429, 1909. 



180 ON MAMMALS FROM WESTERN CHINA. 

It is with great pleasui^e that I name a Chinese animal aftei' 
Pere David, the famous French naturaKst and collector, to whom 
Science owes the discovery of the wonderful series of animals 
on wdiich Milne-Edwards's ' Reeherches Mammiferes ' was based. 
Kuatun, the locality of Rhizomys davidi, was itself discovered as 
:i collecting-station by Pere David, though we owe its more recent 
exploitation to our own countrymen. 

43. Lepus sechuenensis de Wint. 

d. 2258, 2261. $. 2259. 30 miles N.W. of Tau-chow, 
Kan-su. 11,000'. 

Although described as from Sze-chwan, it is probable that this 
species really belongs to the fauna N. and N.W. of the Tsin-ling 
range. The type was said to be from NoHh-viestern Sze-chwan, 
and that part of the Province extends into the faunistic region 
of which Kan-su is a part. No Hare is as yet known from the 
typical forested area of Western Sze-chwan. 

44. OcHOTONA CANSA Lyon. 

5 . 2236, 2243, 2248, 2249. ? . 2226, 2244, 2247. Mts. S.E. 
of Tau-chow, Kan-su. 

Tau-chow (Tao-eheo) is the type locality of Dr. Lyon's species. 

45. OcHOTONA TIBETANA, M.-Edw. 

6 . 2325. $ . 2324. 23 miles S.E. of Ta-tsien-lu, Sze-chwan. 
10,000'. 

c^. 2387. Ta-tsien-lu. 11,600'. 
d. 2525. Omi-san. 9500'. 

These specimens indicate that 0. tihetana has a wide range of 
colour- variation, very much as in the allied 0. hodgsoni. 

46. Capreolus bedfordi Thos. 

S . 2203, 2205, 2207, 2210. 9 . 2204, 2208, 2209. 60 miles 
S.E. of MiH-chow, Kan-su. 8000'. 

47. MOSCHUS SIFANICUS Blichn. 

2 (immature). 2223. 10 miles S. of Tau-chow, Kan-su. 8500'. 
The first example of this striking species that has come to the 
British Museum. 

48. Gazella sp. 

cJ (young). 2260. 30 miles N.W. of Tau-chow, Kan-su. 11,000'. 



No. 87. 

ABSTRACT OF THE PROCEEDINGS 

OF THE 

ZOOLOGICAL SOCIETY OF LONDON.* 

Novemljer 15th, 1910. 

Dr. S. F. Haemer, M.A., F.R.S., Vice-President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a Report on the additions that had been 
made to the Society's Menagerie during the months of June, 
July, August, September, and October 1910, 

Mr. J. Lewis Bonhote, M,A., F.L.S., F.Z.S., exhibited a spirit- 
specimen of a young Cairo Spiny Mouse [Acomys cahirinus), 
about twelve hours old, pointing out the advanced state of the 
young at birth as compared with the common House-MoTise, and 
remarked that the period of gestation in Acomys was eleven days 
at the most, and that the usual number in a litter was three. 

Mr. Bonhote also remarked on a pair of hybrids between the 
Bramble- Finch [Fringilla mo7itifr mgilkc) s.nd the Chaffinch (Frin- 
gilla coelebs), which he had deposited at the Gardens and which 
represented a cross bred for the first time in 1907. He drew 
attention to the resemblances in the plumage of these birds, and 
stated that the fertility of the hybrids had not yet been tested. 

Mrs. R. Haig Thomas, F.Z.S., exhibited a series of skins 
illustrating an experiment in Pheasant-breeding, and gave an 
account of the production of Thaumalea ohsciira in the F, 
generation from a cross between 1\ amhersti $ and T. j^icta S • 

Dr. W. NicoLL and Prof. E. A. Minchin, M.A., Y.P.Z.S., exhi- 
bited specimens of two species of Cysticercoids found in the body- 

* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W,, on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it raay be obtained on tne 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in adva iice. 



42 

cavity of rat-fieas {Ceratophyllus fasciatus). One of these species 
]iad been found in about foui' per cent, of the fleas examined and 
had been shown, by means of feeding experiments carried on by 
Dr. Nicoll, to be the larval form of the common rat-tapeworm 
{Ilymenolepis diminuta). The other species, of which so far only a 
single example had been found, was probably the larva of Hyme- 
nolepis onurina, a species occurring in rats and mice, arid very 
similar to, possibly identical with, H. nana, a dangerous tapeworm 
of man. The chief interest of the discovery lay in the fact that 
no intermediate host was previously known for this tapeworm. 

The Hon. IST. Charles Rothschild, M.A., F.Z.S., exhibited 
sevei'al species of Fleas which were of interest in connection with 
the spreading of plague by these insects. 

Mr. R. I. PococK, F.L.S., F.Z.S., exhibited a living specimen 
of the Black Rat {Mihs rattus) from Sark, which had recently 
been presented to the Society's Mena.gerie by Mrs. C, Russell. 

Mr. R. I. PococK also exhibited a female hybrid, bred in the 
Society's Gardens, between a male Black Lemur {Lemur macaco) 
and a female of the Red-fronted variety of the Fulvous Lemur 
(Lemur fulvus rujifrons), and pointed out that the offspring re- 
sembled neither of its parents. It had not inherited the facial 
fringe of its father nor the white over the eyes and on the fore- 
head of its mother, this area of the head being dark ashy black 
with a deeper tinted central line, more approaching the colour 
seen in the Black-fronted variety of the Fulvous Lemur (L. fulvus 
nigrifrons). 

Mr. J. Lewis Bo]S[hote, M.A., F.L.S., F.Z.S., read a paper 
dealing with some experiments he had made on the occurrence of 
the webfoot character in Pigeons. After referring to Mr. R. 
Staples Browne's paper on the subject in the P. Z. S. for 1905, in 
which that gentleman had shown the webfoot to be a simple 
Mendelian recessive, Mr. Bonhote instanced further cases from 
the lofts of Mr. F. W. Smalley, F.Z.S., that bore out Mr. Staples 
Browne's conclusions. Both these gentlemen, however, had been 
kind enough to give the author birds from their strains, and in 
the first instance when webbed bii'ds from the different strains 
were crossed an irregular result — namely, 4 normal and 1 webbed 
— was obtained. Matings from these birds were continued, and 
the results were, in almost every case, contrary to Mendelian 
expectations, normals throwing webs, and webs throwing normals. 
After discussing various suggestions, Mr. Bonhote came to the 
conclusion that no really satisfactory explanation was forthcoming. 
The Mendelian inheritance was apparently there, but dominated 
and modified b}' some other agency, and he had been able to find 
no single explanation which would cover all the results. 



43 

Mr. Edward Degex, F,Z.S., read a papei^ entitled " Notes on 
the little-known Lizard Lacerta jacksoni Blgr., with special re- 
ference to its Cranial Characters," based on a series of specimens 
recently procured by Mr. R. Kemp in British East Africa. 

Mr. G. A. BouLENGER, F.R.S., V.P.Z.S., read a paper " On 
Lacerta peloponnesiaca Bibr.," which contained a new description 
of this little-known lizard, made from living specimens in the 
Society's Gardens, with a view to fixing its correct position in the 
genus Lacerta. 

A paper entitled " Remarks on two Species of Fishes of the 
genus Gohius, from observations made at Roscoff," was read by 
Mr. E. G. Boulenger, communicated by Mr. G. A. Boulenger, 
F.R.S., Y.P.Z.S., dealing with the specific distinction of Gohiios 
^linutus and &. mia'ops. 



The next Meeting of the Society for Scientific Business will 
be held on Tuesday, November 29th, 1910, at half-past Eight 
o'clock P.M., when the following communications will be 
made : — 

1. Dr, H, B. Fantham, B.A., F.Z.S., and H, Hammond Smith, 

M.R.C.S., L.R.C.P., F.Z.s ! 

On a Possible Cause of Pneumo-enteritis in the Red Grouse 
(Lagopus scoticus). 

2. F. E. Beddard, M.A., F.R.S., F.Z.S. 

On the Alimentary Tract of certain Birds, and on the 
Mesenteric Relations of the Intestinal Loops. 

3. Prof. A. Cabrera, C.M.Z.S. 

On the Specimens of Spotted Hyaenas in the Biitish Museum 
(Natural History). 

4. Dr. J. F. Gemmill, M.A., D.S c. 

The Development of Solaster enchoa Forbes. 



44 

Tke following communications have been received : — . 

1. George P. Farean. 

Plankton from Christmas Island, Indian Ocean. — I, On 
Copepoda of the Family Ooiycseidse. 

2. W. N. F. WOODLAKD, F.Z.S. 



On the Structure and Function of the Gas-Glands and Retia 
Mirabilia associated with the Gas-Bladder of some Teleostean 
Fishes, with notes on the Teleost Pancreas. 

3. H. R. Hogg, M.A., F.Z.S . 

On some New Zealand Spiders. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 

Zoological Society op London, 
Regent's Park, London, N.W. 
Nouemler 22nd, 1910. 



MOTOR eUSES FOR THE SCIENTIFIC MEETINGS. 

On the night of each Scientific Meeting Motor 'Buses 
will run direct to the Gardens from Regent's Park Tube 
Station, starting at 8.5 and 8.20 p.m. The return 
'buses after the Meeting will leave the Gardens at 10.15 
and 10.35 p.m. These 'buses will be labelled m 

PRIVATE and no fare will be charged to Fellows 
attending the Meetings. 



No. 88. 

ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON* 

Noveml)er 29tli, 1910. 



Dr. Henry Woodward, F.R.S., Vice-President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

Mr. D. Seth-Smith, F.Z.S., Curator of Birds, exhibited, through 
the kindness of Mons. Pauvvels, a blue variety of the Budgerigar 
(Melopsittacus undulatus), as well as a yellow variety and a 
normal green specimen. 

The two varieties showed distinctly the splitting up of the 
component coloration, the blue bird lacking all trace of yellow 
pigment, while in the yellow variety the blue colouring of the 
normal bird had disappeared. 

Dr. William Nicoll, of the Lister Institute of Preventive 
Medicine, gave a demonstration of his method for the collection 
of Trematodes. 

Dr. R. T. Leiper, F.Z.S., exhibited two photographs and 
some specimens showing Onchocerciasis in Beef imported from 
Queensland. 

Dr. H. B. Fantham, F.Z.S., and Dr. H. Hammond Smith, 
F.Z.S., contributed a paper " On a Possible Cause of Pneumo- 
enteritis in the Red Grouse {Lagopus scoticus)." The authors 
recorded that in Grouse-chicks dying of Coccidiosis, many of 
which showed symptoms of pneumonia, they found Coccidian 
oocysts in the bronchioles, bronchi, and trachea. The Coccidian 
cysts in the bronchioles were probably capable of setting up 



* This Abstract is published by the Society at its offices, Zoological Gardens, 
Eegent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



46 

sufficient irritation to account for the pneunaonic symptoms. 
These observations were interesting as showing that the much 
criticised views of Klein, Tegetmeier, and others on " pneumo- 
enteritis " as a cause of mortality in Grouse may have some 
foundation in fact. 

Dr. J. F. Gemmill, M.A., D.Sc, Lecturer on Embryology in the 
University of Glasgow, gave an account, illustrated by lantern- 
slides and specimens, of his memoir on " The Development of 
Solaster encleca Forbes," communicated to the Society by Prof. 
J. Arthur Thomson, F.Z.S. 

The author described the ovaries and ova and the processes 
of spawning, fertilization, segmenta,tion, and gastrulation, and then 
dealt with the characters of the free-swimming larvse and the 
changes related to the metamorphosis. He discussed the develop- 
ment of the internal cavities and of the skeleton, and described 
the methods he had employed in obtaining and rearing the larvae. 

The memoir, in addition to details of adult anatomy, contained 
a description of the following points in development : — 

Segmentation total equal ; blastula by egression ; gastrula by 
invagination ; early closure of blastopore ; no larval mouth. 

Free-swimming larva with three arms and a muscular sucker ; 
attachment by sucker. 

Metamorphosis such that while in point of external form the 
left side of the larva becomes the oral surface of the starfish, in 
reality the epiderm of the oral surface is derived in great part fi'om 
that of the anterior part of the early bilateral larva, and conversely 
the aboral epiderm is derived chiefly from that of the posterior end 
of the larva. 

Archenteron dividing into anterior and posterior vesicles and 
middle chamber or enteron. Anterior vesicle giving rise to pre- 
oral, axial, and epigastric cceloms, hydroccele, dorsal sac, internal 
oral sinus, and part of external oral sinus. Posterior vesicle 
giving rise to hypogastric and pharyngeal coeloms, rest of external 
oral sinus, aboral circular sinus, and genital rachis. 

The full set of hydroccele pouches completed slowly, the series 
running in the watch-hand direction as seen from the oral side ; 
opening of stone-canal between radii I and II ; anus in inter- 
radius V/YI. 

Terminal plates double ; rest of aboral plates showing atypical 
distribution. 

A larval nervous system and a statolith -like body in the 
posterior ccelom. 






Mr. F. E. Beddard, M.A., F.K.S., F.Z.S., Prosector to the 
Society, presented a paper " On the Alimentary Tract of certain 
Birds, and on the Mesenteric Relations of the Intestinal Loops," 
based on notes he had accumulated relative to the viscera of 
Birds which had died in the Society's Gardens. The paper dealt 



47 

more particularly with species that had not been carefully studied 
from the point of view of the convolutions of the intestine, and 
attention was called to a considerable series of Birds. 

Mr. Oldfield Thomas, F.R.S., F.Z.S., communicated a paper 
by Prof. Angel Cabrera, C.M.Z.S., "On the Specimens of Spotted 
HyjBnas in the British Museum (Nat. Hist,)," in which he de- 
scribed thi-ee apparently new foi-ms. 



The next Meeting of the Society for Scientific Business will 
be held on Tuesday, December 13th, 1910, at half-past Eight 
o'clock P.M., when the following communications will be 
made : — 

1. Zoological yoMEycLATURE. 

Dr. W. E. Hoyle, M.A., F.Z.S., will explain the Report of 
the International Commission on Zoological Nomenclature pre- 
sented to the Graz Meeting of the International Zoological 
Congress, 1910, and will refer in particular to the proposals 
made for the protection of well-known zoological names. 

2. E. S. Goodrich, M.A., F.R.S., F.Z.S . 

On the Segmentation of the Occipital Region of the Head in 
the Batrachia Urodela. 

3. Dr. W. N. F. Woodland, F.Z.S. 

On the Structure and Function of the Gas-Glands and Retia 
Mirabilia associated with the Gas- Bladder of some Teleostean 
Fishes, with notes on the Teleost Pancreas. 

4. Oldfield Thomas, F.R.S., F.Z.S. 



The Mammals of the Tenth Edition of Linnseus : an Attempt 
to fix the Types of the Genera and the exact Bases and Localities 
of the Species. 



The following communications have been received : — 

1. George P. Farran. 

Plankton from Christmas Island, Indian Ocean. — I. On 
Oopepoda of the Family Corycseidse. 

2. H. R. Hogg, M.A., F.Z.S. 

On some Kew Zealand Spidets. 

3. Dr. F. D. Welch, F.Z.S . 

Observations on different Species of Hylohates now or recently 
living in the Society's Gsivdens,a.nd a Symphalangns S7/7idactytus, 
with Notes on Skins in the Natural History Museum. 

4. R. Lydekker. 



On Tragelaphus huxtoni. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary, 

Zoological Society of London, 
Regent's Park, London, N.W. 
December Qth, 1910. 



MOTOR BUSES FOR THE SCIENTIFIC MEETINGS. 

On the night ©f each Scientific Meeting Motor 'Buses 
will run direct to the Gardens from Regent's Park Tube 
Station, starting at 8.5 and 8.20 p.m* The return 
*buses after the Meeting will leave the Gardens at 10.15 
and 10.35 p.m. These 'buses will be labelled 
PRIVATE and no fare will be charged to Fellows 
attending the Meetings. 



No. 89. 



ABSTRACT OF THE PROCEEDINGS 



OP THE 



ZOOLOGICAL SOCIETY OF LONDON.^ 

December 13th, 1910. 

G. A. BouLENGER, Esq., F.R.S., Vice-President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a Report on the additions that had been 
made to the Society's Menagerie during the month of November 
1910. 

Dr. H. Hammond Smith, F.Z.S., exhibited a mounted specimen 
of a male Red Grouse, from Abington, Lanarkshire, which dis- 
played a curious variety of the ordinary plumage of this species. 

Mr. D. Seth-Smtth, F.Z.S., Curator of Birds, exhibited some 
skins of the Australian Yellow-rumped Finch [Munia Jlavi- 
prymnci). These birds had been kept alive in an outdoor aviary 
in England, and had developed certain markings tending towai'ds 
those of another closely allied species, Munia castaneithorax. 
The exhibitor attributed this to the fact that the former species 
was a desert form of the latter, and when placed in a humid 
environment tended to revert to the plumage of the latter. He 
referred to a paper he had published on this subject in the 
' Avicultural Magazine,' 1907, p. 195. 

Mr. Edwin S. Goodrich, M.A., F.R.S., F.Z.S., read a paper 
" On the Segmentation of the Occipital Region of the Head in 
the Batrachia Urodela," based on his studies of the development 



* This Abstract is published by the Society at its offices, Zoological Grardens, 
Eegent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications; but it maybe obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



of the liead region of the Axolotl [Aonhly stoma tigrinum). The 
head of the Axolotl contained three segments behind the auditory 
capsule. Three metaotic somites were developed in these 
segments, of which the first soon disappeared, and the second and 
third contributed to the formation of the temporal muscle. To 
the first segment belonged the glossopharyngeal nerve ; to the 
next two the vagus. The occipital condyles were developed 
betweea the third and fourth somites. The two hypoglossal 
roots corresponded to the fourth and fifth metaotic segments, and 
passed out in front of the first and second vertebi'se. In the 
Amniota all these segments were included in the head. The 
skull of an Amphibian was thus shorter than that of a Mammal, 
yet the condyles were homologous in the two animals. The 
shifting backwards or forwards of , the condyles was brought 
ahout, not by the inter- or excalation of segments, but by a 
transposition from one segment to another. The sliifting of the 
condyles was comparable to the transposition of the limbs on the 
trunk-segments. 

Mr. Oldfield Thomas, F.K.S., F.Z.S., read a paper entitled 
" The Mammals of the Tenth Edition of Linnsexis : an Attempt 
to fix the Types of the Genera and the exact Bases and Localities 
of the Species." It was shown that by the use of tautonymy the 
types of nearly all the Linnean genera could be definitely fixed, 
the conclusions arrived at by this means agreeing in most cases 
with common usage. The type of Simia, however, would not be 
,S. fsatyrus but S. sylvana, and of Dasyjnis D. novemcinctus instead 
of D. sexclnct'us, the consequences of which changes were pointed 
out. 

Fygafhrix, as represented by the two species nemceus and 
nigripes, was shown to be generically distinct from Fresbytis, so 
that the latter name still remained available for the ordinary 
Langurs. 

Changes in specific names, due to a complete examination, 
were shown to be less numerous than might have been expected, 
while the stability of mammalian nomenclature was much 
increased by avoiding the danger of what such an examination 
might lead to. 

Type localities, derived from the original authors quoted by 
Linnseus, were defined for a considerable number of the species. 

Dr.W. E. HoYLE, M.A., F.Z.S., English Member of the Inter- 
national Commission on Zoological ISTomenclature, explained the 
following Report presented to the Graz Meeting of the Inter- 
national Zoological Congress, and referied in particular to the 
proposals made for the protection of well-known zoological 
names. 

A discussion followed on the portion relating to the formation 
of an Oflicial List of most frequently used Zoological Names. The 
feeling of the Meeting was very strongly in favour of the Inter- 



51 

national Congress giving its authority to tlie formation of a List of 
Zoological Names, the significance of which should not be. altei'ed 
by application of the rules of the Intei-national Code. It was 
vmanimously agreed to accept the action of the Congress if it 
would adopt this coui'se. 

Report of the iNTERiifATiONAL Commission on 
ZooLoaiCAL Nomenclature. 

Financial Aid from the Smithsonian Institution. — Owing to 
the amount of clerical work connected with the studies conducted 
by the Commission, it has been found veiy difficult in the past 
for the Commission to render its decisions as promptly as desirable. 
This difficulty has now been overcome by the genei'ous grant of 
the sum of $2700 by the Smithsonian Institution ; said sum is 
available at the discretion of the Commission at any time during 
the three years following the grant. 

In addition, the Smithsonian Institution has placed at the 
disposal of the Commission the siim of iS500 to be used in p;il)- 
lishing the " Opinions " rendered by the Commission in its 
function as a Court of Appeal. An aiTangement has been made 
between the Secretary of the Smithsonian Institution and the 
Secretary of the Commission, whereby the " Opinions " Avill be 
published by the Institution and forwarded to 1100 libraries, to 
the Members of the International Zoological Congress, and to a 
limited list of specialists. 

Opinions rendered. — Since October, 1909, the Commission 
has rendered 23 " Opinions " (Nos. 6-28), which are now in press 
and which will soon be sent to all Members of the Congress. A 
number of cases are still befoi-e the Commission for study and 
will be passed upon in the near future. The svimmaries of 
Opinions 6-28 are as follows : — 

6. In case of a genus A, Linnaus, 1758, with two species Ah and Ac. — When 

a latei" author rlivicles the genus A, species Ah and Ac, leaving genus A 
only species Ah, and genus C, nionotypic, with species Cc : 

The second author is to be consti'ued as having fixed the type of the 
genus A. [See Article oO.] 
Vote : Affirmative 14 ; negative 0; not voting 1. 

7. On the interpretation of the expression "n. g., n. sp." under Article 30 {a). — 

The expression " n. g., n. sp.," used in publication of a new genus for 
which no other species is otherwise designated as genotype, is to be 
accepted as designation under Article 30 (a). 
Vote : Affirmative 8 ; negative 3 ; not voting 2 ; vote both ways 2. 

8. On the reteniion of ii or i in specific patroni/'inic names, under Article 14 (c) 

and Article 19. — Specific patronymics originally published as ending in 
ii (as schrankii, ehheshornii) are, according to Article 19, to be retained 
in their original form, despite the provision of Article 14 (c), which 
provides that they should have been formed with only one i. 
Vote: Afiirnialive 11 ; negative 1 ; not voting 2 ; vole both ways 1. 



52 

9. The use of the name of a composite genus for a component part requiring 

a name. — The decision as to whether the name of a composite genus, 
wheii made vip wholly of older genera, is tenable for a component part 
requiring a name, depends upon a variety of circumstances. There are 
circumstances under which such name may be used, others under which 
it may not be used. (Article 30.) 
Vote; Affirmative 13; negative 0; not voting 2, 

10. Designation of genotypes for genera puhlished with identical limits. — If 

two genera with the same linaits ai-e formed independently by different 
authors, without designation of genotypes, aiiy subsequent author may 
designate the genotypes ( Ai-t. 30 g) ; and if the types designated are n»t 
specifically identical, the two generic names may (other things being 
equal) be used for restricted genera containing the types in question. 
(Article 25.) 
Vote : Affirmative 9 ; negative 4 ; not voting 2. 

11. The designation of genotypes hy Latreille, 1810. — The " Table des genres 
avec I'indication de I'espece qui leur sert de type," in Latreille's (1810) 
' Considerations generales,' should be accepted as designation of types 
of the genera in question. (Article 30.) 

Vote: Affirmative 12 ; negative 1 ; not voting 2. 

12. Stephanoceros fimhriatus (Goldfnss, 1820) vs. Stephanoceros eichkomii 
Ehrenberg, 1832. — The generic name Stephanoceros, 1832, is to be vised 
in preiereuce to Coronella, 1820 (pre-occupied, 1768); the specific name 
fimhriatus, 1820, takes prei^edence o>er eichkomii, 1832, which is ad- 
mittedly (Ehrenberg, 1832 b, 125, and 1838 a, 400-401) yim/w/a^afs, 1820, 
renamed. Ehrenberg was right in rejecting Coronella, 1820, but in 
error in rejecting fimhriatus, 1820; no reason is apparent for per- 
petuating his eiTor. 

Vote : Affirmative 14 ; negative ; not voting 1. 

13. The specific name of the Samd-Crah. — Catesby's (1743) pre-Linnsean name 
arenarius is not available under the Code, although "reprinted" in 
1771; quadrattis, 1793, is stated to be pre-occupied; albicans, 1802, 
being the next specific name in the list becomes valid, under the 
premises submitted. 

Vote: Affirmative 12 ; negative 0; not voting 3. 

14. The type S2)ccies of Etheostoma Eafinesque, 1819. — The designation of 
E. blennioides Eafinesque, 1819, as type oi Etheostoma ^^Oxveaqiie, 1819, 
by Agassiz, 1854, is not invalidated by the fact that Agassiz used as basis 
for his generic diagnosis characters taken from an erroneous specific 
determination of 1839. Not only does Agassiz distinctly state that 
" Eth. blennioides Eaf." is type of " Etheostoma Raf.," but even if the 
question of the erroneous identification of E. blennioides by Kirtland be 
taken into consideration, the conclusion must be drawn that this erroneous 
identification did not exclude the original specimens of E. blennioides 
from being covered by this specifiic name; on the contrary, the name 
as used by Kirtland, 1839, still involved the type specimens; removing 
now the erroneously determined specimens of 1839, which by article 
30 e (a) are excluded from consideration in designating the genotype, 
the original type specimens of 1819 remain and, upon the premises 
submitted, represent the type of the genus. 

Vote : Affirmative 9 ; negative 4 ; not voting 2. 

15. Craspedacusta sowerbii Lankester, 1880, n. g., n. sp., vs. Limnooodium 
victoria AUman, 1880, n. g., n. sp., a freshwater Medusa. — Craspeda- 
custa sowerbii Lankester, 1880, June 17, has clear priority over Liimio- 
codium victoria AUman, 1880, June 24. Presentation of a paper before 



53 

a scientific society does not constitute puLlication in tlie sense of the 
Code. Tlie Commission is without authority to sanction usage in con- 
travention of the provisions of the Code. 

Vote : Affirmative 15 ; negative 0. 

16. The status of prehinominal specific names (pjihlishcd prior to 1758) mider 
Art. ?>0d. — In deciding whether a case of absolute tautonyinyis present 
(under Art. 30 d), the citation of a clear prebinomina.l specific name in 
synonymy is to be construed as complying with tlie demands of 
Art. 30 d. Examples: Eqims cahalh/s (Equus cited in synonymy in the 
sense of "the horse"), Alca torda {Alca cited in synonymy in the 
sense of " the Alca "). 

Vote : Affirmative 10 ; negative 2 ; not voting 3. 

17. Shall the gevera of Weher, 1795, he accepted 1 — Weber's 'Nomenclator 
Entoraologicus,' 1795, complies with the requirements of Article 25 ; 
hence the genera in question are to be accepted, in so far as they 
individually comply with the conditions of the Code. 

Vote : Affirmative 12 ; negative 1 ; not voting 2. 

18. Tlie ti/pe of Tljdriis 8chneider,\1'i9. — On basis of the premises, ms^ims 
Schneider, syn. hydrus Pallas, is type of Hydrus Schneider, Art. 30 d. 

Vote : Affirmative 14 ; negative ; not voting 1. 

19. Flesiops vs. Pharoptcryx. — From the evidence, it is not clear that this 
case is one of nomenclatorial rather than zoological nature. So far as 
the evidence goes, the question as to wliether Eiippell was in error in 
accepting Plesiops as identical with Pharoptcryx must be answered from 
a systematic point of view. If from our present-dny conception of 
generic limits, Riippell was correct, no reason is apparent for not 
accepting his nomenclatorial decision. 

Vote : Affirmative 31 ; negative 1 ; not voting 3. 

20. Shall the genera of Gronow, 1763, he accepted? — Gro-now. 1763, is binary, 
though not consistently binominal. Article 2r> demands that an author 
be binary, and Article 2 demands tliat generic naroes be uninominal. 
Under these Articles, Gronow's genera are to be accepted as complying 
with the conditions prescribed by the Code to render a name available 
under the Code. 

Vote : Affirmative 11 ; negative 1 ; not voting 3. 

21. Shall the gevera of Klein, 1744, reprinted hy Walhaum, 1792, he 
accepted .?— When Walbaum, 1792. reprinted in condensed form (but 
did not accept) the genera of Klein, 1744, he did not thereby give to 
Klein's genera any nomenclatorial status, and Klein's genera do not 
therefore gain availability under the present Code by reason of being 
quoted by Walbaum. 

Vote: Affirmative 12 ; negative 0; not voting 3. 

22. Ceraticthys vs. Cliola. — Whatever Baird's original intentions may have 
been, he and Girard originally published (1853) Ceraticthys as a mono- 
typic genus, describing the genotype (C vigilax) and giving no indica- 
tion that there were any intentions other than to publish a " n. g., 
n. sp." Under Article 30 c, vigilax is the type of Ceraticthys. 

Vote : Affirmative 12; negative 0; not voting 3. 

23. Aspro vs. Cheilodiptertis, or Amhassis.— Under the premises given, 

Centropomus macrodon may be taken as type of Aspro. 1802, and this 
generic name suppressed as synonym of Cheilodip)terus, thus safeguard- 
ing Amhassis. 

Vote : Affirmative 8 ; negative 1 ; not voting 6. 



54 

24. Antonnarius Cominerson, 1791, and Cuvier, 1817, vs. Histrio Fischer, 

1813. — Antennarius Commerson is an uuinoininal generic name (Art. 2) 
of an author who used a binary (Art. 25) (though not binominal) 
nGmenclature. It received nonienclatorial status by virtue of its 
publication by Lacepede, 1798, and should date from tliat time instead 
of fr^m Cuvier, 1817. It is therefore not necessary to suppress it in 
favour of Histrio, 1813. 

Vote : AiErmative 12 ; negative ; not voting 3. 

25. Bamesiella Tornquist, 1899, vs. Bamesella Walcott, 1905. — Under 
Article 36, Eecommendations, it is not necessary to reject Damesella, 
1905, because of the existence of JDamesicUa, 1898 (1899'?). 

Vote : Affirmative 11 ; negative 1 ; not voting 3. 

26. Cypsilurus vs. Cypsehirus. — In view of the number of typographical 
errors in Svs'ainson, 1838 and 1839, the Commission is oi the opinion 
that Cypsilurusis, an evident typographical error and should be corrected 
to Cypselurus. 

Vote : Affix'mative 10 ; negative 1 ; not voting 4. 

27. Ewpfelia and Bupellia vs. Bilppellia. — Since a typograpjiical error is 

evident, Buppelia and Bupellia should be corrected to liujjpetlia. 

Vote : Affirmative 9 ; negative 1 ; not voting 6. 

28. Shall the ' Nouvelle Classification' of Meigen, \SQO, he given precedence over 
Meigen's ^Versuch,' 1803? — The generic names contained in Meigen's 
' Nouvelle Classification,' 1800, must take precedence over those in his 
' Versuch,' 1803, in every case vs^bere the former are found valid under 
the International Code. 

Vote : Affirmative 11 ; negative ; not voting 4. 

Official List of most frequently used Zoological Xames. — 
Tliere is a clesii^e on the part of some zoologists that certain 
veiy commonly used zoological names should be excej^ted from 
the application of the LotW of Priority, and a proposition to this 
effect has been presented to the Commission from the British 
Association for the Advancement of Science and the Eastern 
Branch of the American Society of Zoologists. That this desire 
is so widespread and so deeply rooted as is assumed by some of 
our colleagues has not been confirmed by inquiries made by 
several members of the Commission. Further, an effort made by 
the Secretary to collect from zoologists the most commonly used 
and most important generic names has as yet met with such poor 
success, that the conclusion does not seem entirely unjustified 
that some of our colleagues who may be in favovir of such a list 
are not as yet sufficiently enthusiastic over the proposition to 
induce them to demonstrate their desire by placing into the 
hands of the Commission the data upon which such a list must 
of necessity be based. Further, there are many colleagues who 
are known to us to be directly and enthusiastically opposed to 
such list. 

After careful consideration of the subject and of the many 
difficulties involved, the Commission has decided to propose 
to the Congress the trial of a proposition which it is hoped 



will meet with the nppi^oval of both sides of the controversy, 
namely : — 

(1) The Commission invites all zo (legists to send to the Secretary of the 

CoiDiuission, prior to November 1, I'JIO, a li.st. of lOU zooloo-ical generic 
names vj-hich they consider should be studied in conneciion with the 
preparation of an "Official List." Each name should be accompanied 
either by the name of the author of the generic name, or by an indica- 
tion of the group to which it belongs. 

(2) All systematists are invited to send a separate list of the 50 to 100 

genei'ic names in their specialty which they look upon as the most im- 
portant and most generally used. Each name should be accompanied 
by the full and complete original bibliographic reference, by the name 
of the type species, determined according to Art. 30 of the Inter- 
national Rules, and by the name of the order and family to which the 
genus belongs. 

(3) All zoologists and palseontologists who give courses in General Zoology 

are invited to supply the Secretary with a list of the text-books 
used in said courses, so that said books may be indexed for generic 
names. 

(4) The Commission will alphabetize all the generic names sent in and will 

endeavour, according to circumstances, to determine which are the 100 
to 500 most commonly quoted genera. 

(5) The genera selected will be submitted to specialists in the groups in 

question, who will be requested to submit opinions on the nomenclatorial 
status of said names. 

(6) Upon return of the lists from the specialists, the Commission will 

endeavour to test the names, according to the International Rules, and 
if feasible will publish a list of the genera in question with their most 
commonly used names and their coriect names. 

(7) If the undertaking is successful, the zoologists of the world will be in- 

vited to give to the Commission the benefit of their criticisms not later 
than July 1, 1912, so that the Commission can restudy the names and 
submit to the next Congress : 

(8) An official list of generic names, with their genotypes, and with the 

(9) Proposition that the Congress adopt said list, and a 

(10) Resolution to the effect that no zoologist shall upon nomknclatoriaTj 
grounds change any name in said list itnless he first submits to the 
Connnission his reasons for making the change and unless the Com- 
mission considers the reasons valid. 

The Commission believes that this proposition is feasible, but 
for the present views it in the light of an experiment, dependent 
to no small extent upon the question whether a proper amount 
of co-operation is forthcoming. In this connection the Commission 
takes the liberty of inviting attention to the fact that the great 
advances in nomenclature have been made by colleagues who have 
showed a conviction in their view sufficient to induce them to 
devote some time to the subject. 



56 

Amendments to the " Regies internationales de la Nomenclature 
zoologique." — In its executive sessions the Commission has eon- 
sidered 30 propositions which have been submitted as amendments 
to the present International Rules. Of these propositions, the 
Commission unanimously recommends to the Congress the 
adoption of the following : — 

Art. 4. For the word root, substitute the word stejn. 

Art. 27 {b). For the word larva, substitute the words any stage in the life- 
history. 

Art, 35. Insert as a third paragraph the following : — 

" Specific names of the same origin and meaning shall be considered 
homonyms if they are distinguished from each other only by the following 
differences ; 

(a) The use of ae, oe, and e, as caeruleus, coeruleus, ceruletis ; ei, i, and y, as 

chiropus, cAeiropus ; c and k, as microdon, mikrodon. 

(b) The aspiration or non-aspiration of a consonant, as oxyryncus, oxy- 

rhynclms. 

(c) The presence or absence of a c before t, as autumnalis, auctumnalis. 

(d) By a single or double consonant : litoralis, littoralis. 

(e) By the endings -ensis and -iensis to a geographical name, as timorensis, 

timoriensis." 

Art. 36. Omit from the examples — Maorodon, Microdon; caeruleus, coeruleus, 

ceruleus ; silvestris, sylvestris, silvaticus, sylvaticus ; littoralis, litoralis ; autum- 
nalis, auctumnalis ; dania, damma. 

Appendix F. In the English and German texts, substitute the words trans- 
literation and transliterated for transcription and transcribed, 

AppeiKlix G. In all the three texts, substitute paragraph for rules, and omit 
from the heading in French text the words Regies de la. 

Italian Translation, — The Commission has voted to issue an 
official Italian edition of the International Rules. 



(Signed) CH. WARDELL STILES, 

Secretary of Commission. 



The next Meeting of the Society for Scientific Business will 
be held on Tuesday, February 7th, 1911, at half-past Eight 
o'clock P.M., Avhen the following communications will be 
made : — 

1. Dr. W. y. F. Woodland, F.Z.S. 

On the Structure and Function of the Gas-Glands and Retia 
Mirabilia associated with the Gas-Bladder of some Teleostean 
Fishes, -with notes on the Teleost Pancreas. 

2. Prof. J. CossAR EwART, M.D., F.R.S., F.Z.S. 

Skulls of Oxen from the Roman Militaiy Station at New- 
stead, Melrose. 

3. George P. Farran. 



Plankton from Christmas Island, Indian Ocean. — I. On 
Copepoda of the Family Corycosidpe. 

4. H. R. Hogg, M.A., F.Z.S. 



On some New Zealand Spiders. 



The following communications have been received : — 

1. Dr. F.D.Welch, F.Z.S. 

Observations on different Gibbons of the Genus Hylohates 
now or recently living in the Society's Gardens, and on a 
Symphalangus syndactylas, with Notes on Skins in the Natural 
History Museum. 

2. R. Lydekker. 

On Tragelaphxis huxtmii, an Antelope from Abyssinia, 

,3. Edward G. Boulenger. 

A Contribution to the Study of the Variations of the Common 
Salamander (S'cdamandra maculosa). 

4. G. A. BouLEXGER, F.R.S., Y.P.Z.S. 

On a Collection of Fishes from the Lake Ngami Basin, 
Bechuanaland. 



58 

Comnivmications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Sp.creMtrij 
Zoological Society op Loxdon, 
Regent's Park, London, N.W, 
December 20ih, 1910. 



MOTOR BUSES FOR THE SCIENTIFIC RIEETINGS. 

On the night of each Scientific Meeting Motor 'Buses 

I will run direct to the Gardens from Regent's Park Tube 

[Station, starting at 8.5 and d.20 p.m. The return 

'buses after the Meeting will leave the Gardens at 10.15 

and 10.35 p.m. These 'buses will be labelled 

j PRIVATE and no fare will be charged to Fellows 

attending the Meetings. 



No. 90. 

ABSTRACT OF THE PROCEEDINGS 



OP THE 



ZOOLOGICAL SOCIETY OF LONDON.^ 

February 7th, 1911. 

Prof. E. A. MixcHiN, MA., Vice-President, 
ill the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a Report on the additions that liad been 
made to the Society's Menagerie daring the month of December 
1910. 

Dr. C, VV. Andrews, F.R.S., F.Z.S., exhibited a skull of a 
Sabre-toothed Tiger (Smilodon californicus) from an asphalt 
deposit in California, and pointed to anatomical characters which 
tended to prove that the animal used its large canines for stabbing 
and tearing, not for biting. 

The Secretart exhibited a mounted specimen of the Platypus, 
which had been lent for the purpose by Mr. P. St. Michael 
Podmore, F.Z.S. 

Mr. Edward Gerrard exhibited the head of a Caribou, shot 
by Sir John Rogers, K.O.M.G., in British Columbia, which had 
a distinct third antler over the centre of the orbital arch of the 
frontal bone. There was a slight burr on a level with the skin, 
but no pedicle. It had the appearance as if it could be shed in 
the same manner as the normal antlers. 

A very fine Eland head, obtained by Major Gordon on the 
Bahr-el-G hazel, and three fine heads of White-tailed Deer were 
also exhibited, one pair of antlers being abnormal. 

* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, ii' desired, sent post-free for 
the sum of SLt Shillings per annum, payable in advance. 



Dr. W. N. F. Woodland, F.Z.S., gave an account of a paper 
on the structure and function of the gas-producing mechanism 
(" red body ") found in connection with the gas-bladders of anany 
Teleostei (Physoclisti and Physostomi), After summai-izing some 
of the principal facts known concerning these subjects, the author 
discussed various theoiies already advanced to account for the 
details of gas-production, and showed that the most satisfactory 
hypothesis was a combination of the views of Jaeger and of 
Nusbaum a,nd Reis, supplemented by additional facts and sug- 
gestions then advanced in the paper. Among the additional 
physiological facts adduced were the sti-iking distribution of the 
capillaries of the rete mirahile of the erythrocytolytic granules 
resulting from the action of Jaeger's toxin on the blood, and the 
peculiar endothelium of the arterioles towards the proximal pole 
of the rete. Both of these facts, together with the otherwise 
meaningless conformation of the rete, supj)orted the view already 
advanced by the author that the rete mirahile duplex is essen- 
tially a device to ensure the diffusion into the artei'ioles of the 
toxin poured into the blood by the gas-gland, so that . erythi'o- 
cytolysis shall be well advanced (and therefore the abstraction of 
oxygen made easy) by the time the arterial blood reaches the 
gas-" secreting " cells. The author also made suggestions con- 
cerning the exact meaning of htemolysis in connection with the 
abstraction of oxygen from the blood by the gas-gland cells. The 
first part of the pa.per dealt with the comparative morphology 
of gas-glands. 

Prof. J. CossAR EwART, M.A., F.R.S., F.Z.S., gave an account 
of his memoir entitled "Skulls of Oxen from the Roman Station 
at Newstead, Melrose," illustrating his remarks by lantern-slides. 
He stated that examination of the skulls from Newstead lent 
support neither to the descent of all European cattle from the 
Urus {Bos prhnigenius) nor to the descent of all European, 
Indian, and African breeds from the Asiatic Urus {B. nomadicus). 
He dealt with the evidence to be derived from the maxillse, the 
occiput and the temporal fossse, and stated his conclusions as 
follows : — - 

1. That the Celtic Shorthorn [Bos longifrons) is probably more 

intimately related to the Zebu of India [Bos indicus) than 
to the European Urus [Bos ])rimi genius). 

2. That long premaxillje are usually correlated with an occiput 

of the Bos primi genius type, while short premaxillse are 
usually correlated with an occiput of the Bos acutifrons 
type. 
.3. That polled black Galloway cattle and polled white " wild " 
Oadzow cattle are intimately related to the Urus, that 
fiat-polled Aberdeen- Angus cattle probably include amongst 
their ancestors an ancient Oriental race now represented 
by, amongst others, a Syrian breed with rudimentary 
horns, and that round-polled cattle may belong to a still 
more ancient Oriental race descended from Bos acittifrons 
of the Punjab Siwaliks. 



IVIr. G. P. Farran presented a paper, communicated by Dr. W. 
T. Calman, F.Z.S., on Copepoda of the family CorycseidjB col- 
lected by Sir John Murray, K.O.B., F.Il.S., and Dr. 0. W. 
Andrews, F.R.S., F.Z.S., at Christmas Island. The collection, 
though small in bulk, was exceedingly rich in species, and the 
genus Corycoius was especially well represented. 

A new genus was proposed and sevei'al new species were 
described and figured, 

Mr. H. R. Hogg, M.A., F.Z.S., read a paper on " Some New 
Zealand Spiders," based on a small collection sent by Prof. Charles 
Chilton, of Christchurch, New Zealand. Twelve species and eleven 
genera were represented in the collection, and a new local variety 
of Tetragnatha ferox and four new species were described. 

Mr. Oldfield Thomas, F.R.S., F.Z.S., read a paper on Mam- 
mals collected in the Provinces of Kan-su and Sze-chwan, Western 
China, by Mr. Malcolm Anderson, for the Duke of Bedford's 
Exploration of Eastern Asia. This collection, from a region 
hitherto ahnost unrepresented in the British Museum, was perhaps 
the finest that had ever come from China, at least so far as small 
mammals were concerned. 47 species were included, i-epresented 
by 350 specimens, presented, as on previous occasions, to the 
National Museum by His Grace. 

Besides examples of a number of rare and important species 
not hitherto in the Museum, specimens of the following new 
forms were included : — 

PlECOTIJS ARIEL, sp. n. 

Allied to P. wardi, but much darker in colour. 
Forearm 44 mm. ; thumb, without metacarpal, 9 "2. Skull 
17-2 X 9-4. 

Hah. Ta-tsien-lu, Sze-cLwan. T'ype. Female. No. 2343. 

Myotis altarium, sp. n. 

Size large. Ears long and I'ather narrow. Skull with vei-y 
short muzzle. 

Forearm 45 mm. ; ear 22. Skull 15-2. 

Hah. Omi-san, Sze-chwan. Tyjie. Female. No. 2423. 

SOREX BEDrORDI^, Sp. n. 

A small dark species, with a blackish line down the back. 
Head and body 55 mm.; tail 55 ; hind foot 13. Skull 17*4 x 8-4. 
Hah. Omi-san, Sze-chwan. Tyj^e. Male. No. 2541. 

SOREX WARDI, sp. n. 

Paler than S. hedfordice ; tail bicolor ; brain-case much smaller. 
Head and body 53 mm. ; tail 49 ; hind foot 12. Skull 17 X 8-2. 
Hah. Tau-cho\^^ Kansu. Type. Male. No. 2230. 



SORICULUS SACRATUS, Sp, n. 

Allied to S. caudatus, but brain-case smaller. 

Head and body 60 mm. ; tail 54 ; hind foot 14. Skull 18'1 X 9-5, 

Hab. Omi-san. T^/pe. Female. No. 2485. 

Ohodsigoa smithii, sp. n. 

Proportions of tail as in Ch. hypsihia, but size much larger. 
Head and body 85 mm. ; tail 68 ; hind foot 18. Skull 22*5 x 
10-5. 

Bob. Ta-tsien-lu. Ttjpe. Male. No. 2349. 

MUSTELA russelliana, sp. n. 

Size excessively small. Colour nearly as in M. kathiah. Tail 
not tufted. 

Head and body 133 mm. ; tail 54 ; hind foot 22. Skull 29-3 X 
15'2. 

Hah. Ta-tsien-lu. Type. Female. No. 2388. 

Epimys excelsior, sp. n. 

Allied to E. conftociamcs, but larger. Skull longer and 
slenderer. 

Head and body 178 mm. ; tail 193 ; hind foot 30. Skull 39. 
Upper molars 6*8. 

Rab. Ta-tsien-lu. Type. Male. No. 2379. 

Epimys andersoni, sp. n. 

Still larger than JiJ. excelsior ; tail very long. 
Head and body 164 mm. ; tail 248 ; hind foot 37. Skull 43. 
Molars 8-2. 

Hah. Omi-san, Sze-chwan. Type. Female. No. 2478. 

Apodemus fergussoni, sp. n. 

General characters of A . agrarius, but no trace of a dorsal line. 
Head and body 107 mm.; tail 98; hind foot 24; ear 15. 
Skull 27-7. 

Hab. "Wen-hsien County, S. Kansu, Type. Female. No. 2296. 

Proedromys bedfordi, g. &, sp. nn. (Microtinse). 

Incisors grooved. M^ with posterior lobe cylindrical, external. 
Fur very long. 

Head and body 103 mm. ; tail 41 ; hind foot 18. Skull 26 x 16. 
Hab. S.E. of Min-chow, Kansu. Ty2}e. Female. No. 2214. 

MiCROTUS (CaRYOMYS) EVA, sp. n. 

(Caryomys, subg. n. — Type, M. inez Thos.) 
Allied to M. mez, but tail very much longer. 
Head and body 88 mm. ; tail 50 ; hind foot 16-5. Skull 22-6 x 
13-5. 

Hab. Near Tau-chow, Kansu. 2'y2ye. Male. No. 2238. 



MiCROTUS MALCOLMI, Sp. n. 

A medium-sized species with high arched skulL 

Head and body 103 mm. ; tail 40 ; hind foot 17. Skull 26-2 x 15. 

Hah. S.E. of Tau-chow, Kansu. Type. Male. No. 2245. 

MiOROTUS IRENE, Sp. n. 

Skull flattened ; brain-case small in proportion to muzzle. 
Mj with only four closed spaces. 

Head and body 98 mm.; tail 37; hind foot 18. Skull 24-2 x 
14-8. 

Hah. Ta-tsien-lu, Sze-chwan. Type. Female. No. 2396. 

Myospalax cansus shenseius, subsp. n. 

Colour warmer than in true cansus. Tooth-row decidedly 
longer. 

Head and body 178 mm.; tail 54; hind foot 30. Upper 
tooth-row (alveoli) 11 "9. 

Hah. Yu-lin-fu, Shensi. Type. Male. B.M. No. 9.1.1,216. 

Rhizomys davidi, sp. n. 

Soft fur and short tail of li. vesUtus, but skull scarcely larger 
than in R. siiiensis. 

Head and body 350 mm. ; tail 69 ; hind foot 42. Skull 66 x 
49' 5. 

Hah. Kuatun, Fo-kien. Type. Female. B.M. No. 96.12.1.6. 



The next Meeting of the Society for Scientific Business will 
be held on Tuesday, February 21st, 1911, at half- past Eight 
o'clock P.M., when the following communications will be 
made : — 

1. Dr. H. G. Plimmer, F.R.S., F.Z.S., Pres.R.M.S . 

Report on the Deaths which occurred in the Zoological 
Gardens during 1910. 

2. R. Lydekkbr. 

On Tragelaphus buxtonl, an Antelope from Abyssinia. 

3. Edward G. Boulenger . 

A Contribution to the Study of the Variations of the Common 
Salamander {■Salainaiidra laacaloaa). 



6 

4. G. A. EouLEyGER, F.R.S., V.P.Z.S . 

On a Collection of Fishes from the Lake- ISTgami Basin, 
Bechuanaland, 

5. Dr. F. D. Welch, F.Z.S. 

Observations on different Gibbons of the Genus Hylohates 
now or recently Living in the Society's Gardens, and on a 
Symphalangus syndactytus, with Notes on Skins in the Natural 
History Museum. 



The following communications have been received : — 

1. R. Lydekker . 

{a) A rare Beaked Whale. 
. (6) Age Phases of the Rorqual. 

2. Mrs. E. W. Sextox. 

On the Amphipod Genus Leptocheirus. 
' 3. F. E. Beddard, M.A., F.R.S.. F.Z.S. 

{a) Contributions to the Anatomy of the Anura. — I. Some 
Anatomical Notes upon the Frog Megalophrys (Leptobrachium) 
fern. 

{b) On the Spermatophores in Earthworms of the Genus 
Pheretima {=Ferichceta). 

4. William Nicoll, M.A., D.Sc, M.ll . 

On Three new Trematodes from Reptiles. 

5. G. Stewardsox Brady, M.D., LL.D., D.Sc, F.R.S., C.M.Z.S. 

Notes on Marine Ostracoda from Madeira. 

6. J. Lewis B oxhote, M.A., F.L.S., F.Z.S, and F. W. Smalley , 

F.Z.S. 
On Colour and Colour-pattern Inheritance in Pigeons. 

7. P. Chalmers Mitchell, M.A., D.Sc, Hon.LL.D., F.R.S., 

F.ZS. 

On Longevity and Relative Viability in Mammals and Bird.^ ; 
with a Note on the Theory of Longevity. 



Oommunieations iiitendeil for the .Scientine IVFeetings sliould 
be addressed to 

P. CHALMERS MITCHELL, 

Secretarij. 

Zoological Society of Loxdox, 
Regent's Park, Loxdox, N.VV. 
February \Uh, 1911. 



MOTOR BUSES FOR THE SCIENTIFIC MEETINGS. 

On the night of each Scientific Meeting Motor 'Buses 
will run direct to the Gardens from Regent's Park Tube 
Station, starting at 8.5 and 8.20 p.m. The return 
'buses after the Meeting will leave the Gardens at 10.15 
and 10.35 p.m. These 'buses will be labelled 
PRIVATE and no fare will be charged to Fellows 
attending the Meetings. 



No. 91. 

ABSTRACT OF THE PROCEEDmGS 

OF THE 

ZOOLOGICAL SOCIETY OF LONDON.* 

February 21st, 1911. 



Dr. A. Smith Woobward, F.R.S., Yice- President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a Report on the additions that had been 
made to the Society's Menagerie during the month of January 
1911. 

Mr. A. Heneage Cocks, F.Z.S., exhibited a series of photo- 
graphs of the female Brindled Gnu recently born in the Society's . 
Gardens, and gave a brief account of its growth and coloration. 

Dr. H. Hammond Smith, M.R.C.S., F.Z.S,, exhibited three 
skins of male Pheasants assuming female plumage, sent to him 
by Mr, Arthur Gilbey, and some microscopical specimens of the 
glands prepared by Dr, S. G, Shattock. 

Dr. H. G. Plimmer, F.R.S., F.Z.S., Pathologist to the Society, 
reported on the Deaths which had occurred in the Zoological 
Gardens during the past year, and illustrated his i-emarks with 
a large series of lantern-slides, prepared from pathological material 
obtained in the course of his work at the Gardens. 

Mr. R. Lydekker communicated a short paper on Tragelaplms 
huxtoni, an Antelope obtained by Mr. Ivor Buxton in Abyssinia. 



* This Abstract is published by the Society at its offices. Zoological Gardens, 
Eegent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six SkiUitigs per annum, payable in advance. 



10 

Mr. E. G. BouLENGER read a paper, communicated by Mr. G. A. 
JBouLENGER, F.K.S., V.P.Z.S., on the varieties of the Spotted 
Salamander {Salamandra maculosa), illustrated by lantei-n-slides 
and specimens. One of the principal results of the author's 
study was to lay greater stress on the disposition of the spots 
than on their actual form, size, or colour, and to define two 
principal forms in Central Europe, which had not previou&ly 
been separated with sufficient precision, notwithstanding their 
well-marked geographical distribution. 

The author further dealt with some of the experiments of 
Ur. Kammerer, of Vienna, and the conclusions arrived at by him 
with regard to coloration in relation to environment. 

Mr, G. A. BouLENGER, F.R.S., V.P.Z.S., contributed a paper 
based on a collection of Fishes from the Ijake ISTgami Basin, 
Bechuanaland, made by Mr. R. B. Woosnam, F.Z.S. 

This paper will appear in the ' Transactions ' in due course. 

Dr. F. D. Welch, F.Z.S. , communicated a paper on Gibbons of 
the Genus Hylohates, and on a Siamang Gibbon, recently living 
in the Society's Gardens, with notes on skins in the British 
Museum (Nat. Hist.). 



T\\Q next Meeting of the Society for Scientific Business will 
be held on Tuesday, March 7th, 1911, at half-past Eight 
o'clock P.M., when the following communications will be 
made : — ■ 

Laktern Exhibitioxs :— 

a. Habits of Moose. 

h. Carl Hagenbeck's Tierpark, 

1. The Hon. N. Charles Rothschild, M.A.. F.Z.S. 

Some new Siphonaptera from China. 

2. F. E. Beddard, M.A., F.R.S., F.Z.S . 

{a) Contributions to the Anatomy of the Antu'a. — I. Some 
Anatomical Notes upon the Frog Megaloflirys {Leptobraclimm) 
fea\ 

(6) On the Spermatophores in Earthworms of the Genus 
Pheretiona ( = Ferichceia), 



11 

3. R. Lydekker. 

(«) A rai-e Beaked Whale. 

(6) Age Phases of the Rorqual, 

4. P. Chalmers Mitchell, jNI.A., D.Sc, Hon.LL.D., F.R.S., 

F.Z.S. 

On Longevity and Relative Viability in Mammals and Birds ; 
with a Note on the Theory of Longevity. 



The following communications have been received : — 

1. Mrs. E. AY. Sextok . 

On the Amphipod Genus Leptoolteirus. 

2. William Mcoll, M.A., D.Sc, M.B . 

On Three new Trematodes from Reptiles. 

3. C4. Stewarlson Bhady, M.D., LL.D., D.Sc, F.R.S., C.M.Z.8 . 

Notes on Marine Marine Ostracoda from Madeira. 

4. J. Lewis Poxegie, M.A., F.L.S., F.Z.S., and F. W. Smalley , 

On Colour and Colour-pattern Inheritance in Pigeons. 



Communications intended for the Scientific ]\Ieetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 

Zoological Society of Loxdok, 
Regent's Park, Loxdon, N.W. 
February 2Sth, 1911. 



12 



mOTaR BUSES FOR THE SCiENTIFIG MEETINGS. 

On the night of each Scientific Meetiii{^ Motor 'Buses 
will run direct to the Gardens from Regent's Park Tube 
Station, starting at 8.5 and 8.20 p.m. The return 
*buses after the Meeting will leave the Gardens at 10.30 
and 10.45 p.m. These 'buses will be labelled 
PRIVATE and no fare will be charged to Fellows 



attending the Meetings. 



No. 92. 



ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON; 

March 7th, 1911. 



Dr. A. Smith-Woodward, F.R.S., Vice-President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

Mr. R. I. PococK, F.L.S., F.Z.S., exhibited, on behalf of 
Mr. E. C. Oberholtzer, a large number of lantern-slides and 
photographs illustrating the habits of Moose. Mr. Oberholtzer 
had spent several months in the forested lakeland tributary to 
Rainy Lake, Ontario, photographing and observing the habits of 
Moose, and had presented to the Society the series of enlarge- 
ments exhibited. 

The Secretary exhibited a series of lantern-slides prepared 
from photographs kindly given to liim by Mr. Carl Hagenbeck, 
Silver Medallist of the Society, and illustrating some of the most 
remarkable features of Mr. Hagenbeck's new Tiei-park at Stellingen 
near Hamburg. 

Dr. Karl Jordan, F.E.S., and the Hon. IST. Charles Roth- 
schild, M.A., F.Z.S., F.E.S. , communicated a paper " On some 
Siphonaptera from Northern China." This collection of Fleas 
had been made by Mr. M. P. Andei^son, the Duke of Bedford's 
collector, in the province of Shen-si, and contained altogether 17 
species, of which no less than 13 were new. Some of these 



* This Abstract is published by the Society at its offices, Zoological Gardens, 
Eegent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Siayence, or, if desired, sent post-free for 
tlie sum of Six ShiUinys per annum, payable in advance. 



14 

Avere possibly only geographical developments, but others repre- 
sented distinct types not very nearly allied to any known species. 

Mr. F. E. Beddard, M.A., F.R.S., F.Z.S., Prosector to the 
Society, read a paper dealing with certain points in the anatomy 
of the Frog Megalophrys {Leptobrachiivni) fece, based on specimens 
which had been exhibited in the Society's Gardens. 

Mr. Beddard also read a paper on the Spermatophores in 
Earthworms of the genus Pheretima {^^Perichceta). These 
structures had been found in two species, one of which would be 
described as new, contained in a collection of teri^estrial Oligo- 
chseta from the Philippine Islands, which had been submitted to 
him for examination by the Director of the Scientific Bureau of 
the Philippines. 

Mr. R. Lydekker communicated two short papers entitled 
{a) "A Rare Beaked Whale," and (5) "Age Phases of the 
Rorqual." 

Dr. P. Chalmers Mitchell, M.A., F.R.S., Secretary of the 
Society, gave an account of his memoir entitled " On Longevity and 
relative Viability in Mammals and Biixls ; with a Note on the 
Theory of Longevity." The woi'k was based on a study of the 
records of the duration of life in the Society's Gardens of over 
20,000 individual Mammals and Birds. These were ari-anged 
systematically so as to make possible a comparison of the average 
duration with the maximum duration and what was known or could 
be inferred as to the potential longevity. Such a method gave a 
measure of the effect of the conditions of captivity on the duration 
of life. The memoir discussed some of the results obtained by 
such a comparison, j)articularly with regaixl to the provision of 
artificial heat. In the note on the Theory of Longevity, the 
author briefly reviewed the contributions of Ray Lankester, Weis- 
mann, and Metchnikoff, and stated his conclusion that potential 
longevity was due to constitutional causes, that the constitution 
was adapted to the average specific longevity, and that the corre- 
lation between longevity and reproduction was the reverse of 
what had been suggested by Weismann. 



15 

The next Meeting of the Society for Scientific Business will 
he held on Tuesday, March 21st, 1911, at half-past Eight 
o'clock P.M., when the following commvniications will be 
made : — 

1. Mrs. E. W. Sextox. 



On the Amphipod Genus L&ptocheirus. 

2. J. Lkwis BoxnoTE, M.A.. F.L.S., F.Z.S., and F. W. Smvllev, 
F.Z.S. 

On Colour and Colour-pattern Inheritance in Pigeons. 

8. G. Stewardsox Brady, M.D., LL.D., D.Sc, F.R.S., C.M.Z.S. 
Xotes on Marine Ostracoda from Madeira. 



The folloAving communications have been received : — 

1. William Nicoll, M.A., D.Sc, M.B. 

On Three new Trematodes fiom Reptiles. 

2. J. A. MoRCH. 

On the Natural History of Whalebone Whales. 



Communications intended for the Scientific Meetings shoulil 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 

Zoological Society of Loxdox, 
Regext's Park, Loxdox, N.W, 
March 14^^, 1911. 



16 



MOTOR BUSES FOR THE SCIENTIFIC MEETINGS. 

On the night of each Scientific Meeting Motor 'Buses 
j will run direct to the Gardens from Regent's Park Tube 
Station, starting at 8.5 and 8.20 p.m. The return 
'buses after the Meeting will leave the Gardens at 10.30 
and 10.45 p.m. These 'buses will be labelled 
[PRIVATE and no fare will be charged to Fellows 
j attending the Meetings. 



Exhibitions and Notices (continued). 

Page 
Mr. D. Seth-Smith, F.Z.S. Exhibition of skins of the Austi'alian Yellow-ruinpecl Finch 

{Munia fiavipryvina) 10 1 

Dr. W. E. HoYLK, M. A., F.Z.S. Exphmation of the Eeport of the International Com- 
mission on Zoological Nomenclature presented to the Graz Meeting of the International 
Zoological Congress 101 

PAPERS. 

1. On the Inheritance of the Webi'oot Chai'acter in Pigeons. By J. Lewis Boniiote, 
M.A., F.L.S., F.Z.S. (Text-(ig3. 3 & 4.) 14 

2. Notes on the little-known Lizard Lacerta jacJcsoni Blgr., with Special Eeference to its 

Cranial Characters. By Edward Deoen, F.Z.S. (Text-figs. 5-7.) 11) 

3. On the Peloponnesian Lizard {Lacerta pcloponnesiaca Bibr.). By G. A. Boulenger, 

F.R.S., V.P.Z.S. (Plate I. and Text-fig. 8.) 37 

4. Remarks on Two Species of Fishes of the Genus Gohius, from Observations made at 
Eoscoif. By Edward G. Boulengke 40 

5. On a Possible Cause of Pneumo-enteritis in the Red Grouse {Lagopus scoticus). By 
H. B. Fantham, D.Sc, B.A., F.Z.S., and H. Hammond S.mitii, M.R.C.S., L.R.C.P., 
F.Z.S 4t) 

6. On the Alimentary Tract of certain Birds and on the Mesenteric Relations of the 
Intestinal Loops. By Frank E. Beddard, M.A., F.R.S., F.Z.S., Prosector to the 
Society. (Test-figs. 9-28.) 47 

7. On the Specimens of Spotted Hytenas in the British Museum (Natural History). By 
Prof. Angel Cabrera, C.M.Z.S 93 

y On the Segmentation of the Occipital Region of the Head in the Batrachia Urodela. 
By Edwin S. Goodrich, M.A., F.R.S., F.Z.S., Fellow of Merton College, Oxford. 
(Text-figs. 29-5] .) 1,,1 

9. The Mammals of the Tenth Edition of Linnaeus; an Attempt to fix the Types of the 
Genera and the exact Bases and Localities of tlie Species. By Oldfield Thomas 
F.R.S., F.Z.S ' ' ]oo 

10. The Duke of Bedford's Zoological Exploration of Eastei-u Asia. — XIII. On Mammals 
from the Provinces of Kan-su aiid Sze-chwan, Western China. By Oldfield Thomas 
F.R.S., B\Z.S \ Yoi> 



Plate I. 
Lacerta pel jjionnesiaca pao-e 37 



NOTICE. 

The ' Proceedings ' for tbe jear are issued in four part.s, paged consecutively, 
so that tiie complete reference is now P. Z. S. 1011, p. . . . The Distribution 
is as follows : — 

Part 1 issued in March. 
,, 2 ,, June. 

,, o ,, September. 

,, 4 ,, Decern her. 



' Proceedings/ 1910, Part IV. (pp. 8o7-103;J), were puWislied on 
January 11th, lUU. 



The 'Abstracts of the Proceedings,' Nos. 87 92, 
axe contained in this Part. 






PROCEEDINGS 



GENERAL MEETINaS FOR SCIENTIFIC BUSINESS 



OF THE 



ZOOLOGICAL SOCIETY 

OF LONDON. 



1911. 



PART II. 

CONTAINING PaGES 181 TO 555^ WITH 15 PlATES 

AND 93 Text-figures. 



JUNE 1911. 



PRINTED FOE THE SOCIETY, 
SOLD AT THSm HOUSE IN REGENT'S PARK. 

LONDON : 
MESSRS. LONGMANS, GRBEN, AND CO., 

PATERNOSTER ROW. 



[Price Twelve Shi/iings.] 







ill 



5ti-x> 



'o. \ 



4/.'^ \ ii". ^ ^A 



A 



LIST OF CONTENTS. 
1911, Part II. (pp. 181-555). 



EXHIBITIONS AND NOTICES. 

Page 
Ihe Secretary. Report on the Additions to the Society's Menagerie during the month of 

December 1910 181 

Dr. 0. W. Andrews, F.R.S., F.Z.S. Exhibition of the skull of a Sabre-toothed Tiger 

{Smilodon californicus) 181 

The Secretary. Exhibition of a mounted specimen of the Platypus {Ornithorhynchus 

paradoxus) 181 

:Mr. Edward Gekrard, F.Z.S. Exhibition of the heads of a Caribou {Rangifer tarandus 
caribou), an Bland {Taurotragus derhianus gigas), and three White-tailed Deer 
(JDorcelaphus americanus macrurus), (Text-fig. 52.) 181 

'The Secretary. Report on the Additions to the Society's Menagerie during the month 

of January 1911 314 

Dr. H. Hammond Smith, M.R.O.S., F.Z.S. Exhibition ot skins of male Pheasants assum- 
ing female plumage 314 

Mx. Gr. A. BouLENGER, F.R.S., V.P.Z.S. Notice of a paper " On a Collection of Fishes 

from the Lake Ngami Basin, Bechuanaland " 314 

Mr. Alfred H. Cocks, M.A., F.Z.S. Exhibition of photographs of, and remarks upon, the 

female Brindled Gnu recently born in the Society's Gardens. (Text-figs. 97 & 98.) . . 314 

The Secretary. Exhibition of a series of lantern-slides illustrating some ' of the most 

remarkable features of Mr. Carl Hagenbeck's new Tierpark at Stellingen 358 

Mr. Ernest C. Oberholtzer. Exhibition of lantern-slides and photographs illustrating 

an account of some Observations on Moose 358 

PAPERS. 

11. On the Structure and Function of the Gas Glands and Retia Mirabilia associated with 

the Gas Bladder of some Teleostean Fishes, with Notes on the Teleost Pancreas. By 
W, N. F. Woodland, F.Z.S., The Zoological Department, University College, London. 
(Plates II.-IX. and Text-figs. 53-62.) 183 

12. On Skulls of Oxen from the Roman Military Station at Newstead, Melrose. By J. C, 

EwART, M.D., F.R.S., F.Z.S. (Text-figs. 63-91.) 249 

13. Plankton from Christmas Island, Indian Ocean. — I. On Copepoda of the Family 

Corycffii*d£e. By George P. Farran. (Plates X.-XIV.) 282 

14. On some New Zealand Spiders. By H. R. Hogg, M.A., F.Z.S. (Text-figs. 92-96.) .. 297 

Contents continued on page 3 of Wrapper. 



THE ZOOLOGICAL SOCIETY OF LONDON. 



This Society was fouaded in 182G by Sir Sxajiford Raffles 
Mr. J. Sabine, Mr. N. A. Vigoks, and other eminent Natnralists, 
for the advancement of Zoology and Animal Physiologj', and for the 
introduction of new and curious subjects of the Animal Kingdom, 
and was incorporated by Eoyal Charter in 1829. 



HIS MAJESTY THE KING. 



COUNCIL. 

HIS GEACE THE DUKE OF BEDFORD, KG., Tresident. 



The EaPvL of Altamont, F.S.A. 
Prof. J. "Rose Bradfokd, M.D., 

D.Sc, F.E..S., Vice-President. 
Lt.-Col. Sir E,. Havelock 

Charles, M.D., K.C.V.O. 
Alfred H. Cocks, Esq., M.A. 
The Et. Hon. the Earl of 

Cromer, P.C.,G.C.B., G.C.M.G. 
F. D. Dawtrex Lrewitt, Esq., 

M.A., M.D. 
Charles Drujijiond, Esq,., 

Treasurer. 
Sir Edward Durand, Bt., C.B. 
Frederick Gillett, Esq., Vice- 
President. 
Sidney F. Harmbr, Esq., M.A. 

Sc.D., F.E.S., Vice-President. 



Sir Walter Roper Lawrence, 

Bt., G.C.I.E. 
Sir Edmund G. Loder, Bt. 
E. G. B. Meade-Waldo, Esq., 

Vice-President. 
P. Chalmers Mitchell, Esq., 

M.A., D.Sc, LL.D., F.R.S., 

Secretary, 
W. R. OaiLviE-GRANT, Esq. 
Adrian D. W. Pollock, Esq. 
AuBYN Trevor-Battye, Esq., 

M.A. 

Anthony H. Wingfield, Esq. 
A.Smith AVoodward,Esq.,LL.D., 

F.R.S., Vice-President. 
Henry Woodward, Esq., LL.D., 

F.R.S., Vice-President. 



The Society consists of Fellows, and Honorary, Foreign, and 
Corresponding Members, elected according to the By-Laws. It 
carries out the objects of its foundation by means of the collection 
of living animals, by its Library, and by its Scientific Publications. 

The Office of the Society, where all communications should be 
sent, addressed to "The Secretary," is open from Ten till Five, 
except on Saturday's, when it closes at Two p.m. 

The Library, under the superintendence of Mr. F. H. "Waterhouse, 
is open daily at the above hours, except in September, 

The Meetings of the Society for General Business are held on 
the third Wednesday in every month of the year, except in 
September and October, at half-past Four o'clock p.m. 

The Meetings for Scientific Business are held fortnightly on 
Tuesdays, except in July, August, September, and December and 
January, at half-past Eight o'clock p.m. 

The Anniversary Meeting is held on the 29th, of April, or the 
nearest convenient day, at Four p.m. 

The Gardens are open daily from Nine o'clock until Sunset. 
Mr. K. I, Pocock, F.R.S., F.L.S., is the resident Superintendent and 
Curator of Mammals. Mr. D. Seth-Smith is Curator of Birds 
and Inspector of Works. The Prosectorium for Anatomical and 
Pathological work is under the charge of Mr. Frank E. Beddard, 
M.A., F.E.S., Prosector, assisted by Mr. H. G. Plimmer, F.KS., 
M.li.C.S., Pathologist to the Society. 



TEEMS FOU THE ADMISSION OF FELLOWS. 

Fjjllows pay an Admission Fee of £5, and an Annual Contri- 
bution of £3, due on the 1st. of January, and payable in advance, 
or a Composition of .£45 in lieu thereof; the whole payment, 
including the Admission Fee, being .£50. 

No person can become a Fellow until the Admission Fee and 
first Annual Subscription have been paid, or the annual payments 
have been compounded for. 

Fellows elected after the 31st. of August are not liable for the 
Subscription for the year in which they are elected. 



PEIVILEGES OF PELLOWS. 

Fellows have Personal Admission to the Gardens upon signing 
their names in the book at the entrance gate, and may introduce 
Two Companions daily. 

The Wife or Husband of a Fellow can exercise these privileges 
in the absence of the Fellow. 

Every Fellow is entitled to receive annually 60 undated Green 
Cards, and, when no specific instructions are received, the supply 
will be sent in this form. If preferred, however, 20 Green Cards 
may be exchanged for a book containing 2 Orders for each 
Saturday * throughout the year. A similar book of Sunday Orders 
may also be obtained in lieu of 20 Green Cards. A Green Card 
may also be exchanged for 2 BufE Cards for the use of Children 
under 12 years of age. 

It is particularly requested that Fellows luill sign every Tichet 
before it goes out of their possession. Unsigned Tickets are nob 
available. 

Green and Buff Tickets may be used on any day and in any year, 
but in no case can two Children be admitted with one Adult's 
Ticket, or an Adult be admitted with two Children's Tickets. 

The annual supply of Tickets will be sent to each Fellow on the 
1st. of January in every year, upon filling up and returning the form 
of Standing Order supplied to Fellows. 

Fellows are not allowed to pass in friends on their written 
order or on presentation of their visiting cards. 

Fellows are exempt from payment of the fee for Painting, 
Sketching, and Photographing in the Society's Gardens. 

Fellows have the privilege of receiving the Society's ordinary 
Publications issued during the year upon payment of the additional 
Subscription of One Guinea. This Subscription is due upon the 
1st. of January, and must be paid before the day of the Anniversary 
Meeting, after vrhich the privilege lapses. Fellows are likewise 
entitled to purchase these Publications at 2b per cent, less than 
the price charged to the public. A further reduction of 25 per 
cent, is also made upon all purchases of Publications issued prior 
to 1881, if above the value of Five Pounds. 

Fellows also have the privilege of subscribing to the Annual 
Volume of ' The Zoological liecord,' which gives a list of the Works 
and Publications relating to Zoology in each year, for the sum of 

* The Saturday Orders are not available if the Fellow introduces friends 
personally on that day. 



One Pound Ten Shillings. Separate divisions of volames 39 to 
42 can also be supplied. Full particulars of these publications can 
be had on application to the Secretary. 

Fellows may obtain a Transferable Ivory Ticket admitting 
two persons, available throughout the whole period of Fellowship, 
on payment of Ten Pounds in one sum. A second similar ticket 
may be obtained on payment of a further sum of Twenty Pounds. 

Any Fellow who intends to be absent from the United Kingdom 
during the space of at least one year, may, upon giving to the 
Secretary notice in wanting, have his or her name placed upon the 
" dormant list," and will then be called upon to pay an annual 
subscription of £1 only during such absence, but after three years 
must make a further application to be retained on that list. 

Any Fellow, having paid all fees due to the Society, is at liberty 
to withdraw his or her name upon giving notice in luriting to the 
Secretary. 

Ladies or Gentlemen wishing to become Fellows of the Society 
are requested to communicate with the undersigned. 

P. CHALMEES MITCHELL, 

Secretary. 
Regent's Park, London, N.W., 
June, 1911. 



MEETINGS 

OP THE 

ZOOLOGICAL SOCIETY OF LONDON 

FOE 

SCIENTIFIC BUSINESS. 



1911. 

Ttjesdat, October 24 

November 7 «& 21 



The Cliair will he talcen at Jialf-past Eight odock in the Evenivg 
precisely. 



ZOOLOGICAL SOCIETY OF LONDON. 



THE ZOOLOGICAL RECORD. 

'THE object of the Zoological Kecokd is to give, by means of an 
annual Volume, complete lists of the Works and Publications 
relating to Zoology in all its branches that have appeared during 
the year preceding the issue of the Yolume, together with full 
information as to the points they deal with, arranged in such a 
manner as to serve as an Index to the literature of Zoology in all 
parts of the globe, and thus to form a repertory that will retain its 
value for the Student in future years. 

The ' Zoological Record ' having been amalgamated with the 
International Catalogue of Scientific Literature, Zoology, Volumes 
from 43 onwards can now be obtained only from Messrs. Harrison 
& Sons, except when purchasing complete sets from the Zoological 
Society. 

Under the scheme of amalgamation, Fellows of the Society, and 
Institutions already on the subscription-list, have the privilege of 
subscribing at the old rate of 30.s". per annum, which covers the 
cost of carriage of the volume. The subscription becomes due on 
July Ist. in each year, and lapses if not paid by the 1st. of December 
following. 

The Society is able to supply complete sets of the Record on the 

following terms : — 

Vols. 1 to 37, price £14 10s. net. 

Vols. 38, 39, 40, and 41 at 10s. each net. 

Vol. 42 and onwards at 40s. each. 

The prices for separate volumes are as follows : — 

Vols. 1 to 41 (except Vols. 4 and 6) 10s. each net. 

Vol. 42 at 40s. The price of the 'Zoological Record,' Vol. 43 and 

subsequent volumes, published now by Messrs. Harrison and Co., 

is 40s. each. 

Index Zoologictjs. An alphabetical list of names of genera 
and subgenera proposed for use in Zoology, as recorded in the 
' Zoological Record,' 1880-1900 ; together with other names not 
included in the ' Nomenclator Zoologicus ' of S. H. Scudder. Com- 
piled (for the Zoological Society of London) by Chakles Owen 
Waterhousb and edited by David Shaep, Editor of the ' Zoological 
Record.' London, 1902. Price to Eellows, 18s. ; price to the 
public, 20s., or if sold with a set, 10s. 

Divisions of the ' Zoological Record ' of Vols. 39 to 42 can be 
supplied by the Society, but after Vol. 42 they can be had only of 
Messrs. Harrison & Sons, 46 St. Martin's Lane, "W.C. 

[p. T. o. 



SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD. 

Divisions of the 'Zoological Record,' Vols. 39-42, containing 
the literature of the years 1902-1905, may be obtained separately 
as follows : — 

s. d. 

List of abbreviations of journals, etc 2 net. 

Special Records, viz. : — 

I. General Subjects 2 6 „ 

II. Mammalia 2 6 „ 

IIL Aves 6 „ 

IV. Reptilia and Batrachia 2 6 „ 

V. Pisces 2 6 „ 

VI. Tunicata 1 „ 

VII. Mollusca 4 „ 

VIII. Brachiopoda . . . . 1 „ 

IX. Bryozoa 1 „ 

X. Crustacea 2 6,, 

XI. Arachnida 2 „ 

XII. Myriopoda 1 6 „ 

XIII. Insecta 12 „ 

XIV. Echinoderma 3 6 „ 

XV. Vermes 3 „ 

XVI. Coelenterata 16,, 

XVn. Spongiffi 2 „ 

XVIII. Protozoa 2 „ 

Index of new names of genera and subgenera . 2 „ 

Divisions from Vol. 43 onwards are now supplied by Messrs. 
Harrison & Sons, 46 St. Martin's Lane, London, W^.C. 



P. CHALMERS MITCHELL, 

Secretary. 



Eegbnt's Park, London, N.W. 
June, 1911. 



ZOOLOGICAL SOCIETY OF LONDON. 



LIST OF PUBLICATIONS. 

The scientific publications of the Zoological Society of London 
are of two kinds — " Proceedings/' published in an octavo 
form^ and " Transactions/' in quarto. 

According to the present arrangements^, the " Proceedings" 
contain not only notices of all business transacted at the scien- 
tific meetings, but also all the papers read at snch meetings 
and recommended to be published in the ''Proceedings'^ by 
the Committee of Publication. A large number of coloured 
plates and engravings are issued in the volumes of the 
" Proceedings/' to illustrate the new or otherwise remark- 
able species of animals described therein. Amongst sucli 
illustrations, figures of the new or rare species acquired in a 
living state for the Society's Gardens are often given. 

The " Proceedings " for each year are issued in four parts, 
paged consecutively, on the first of the months of March, 
June, September, and December. From January 1901 they 
have been issued as two half-yearly volumes, indexed 
separately. 

An "Abstract of the Proceedings " is published by the 
Society on the Tuesday following the date of Meeting to 
which it refers. It is issued along with the " Proceedings," 
free of extra charge, to all Fellows who subscribe to the 
Publications, but it may be obtained on the day of publi- 
cation at the price of Sixpence, or, if desired, sent post free 
for the sum of Six Shillings per annum, payable in advance. 
The "Transactions^' contain such of the communications 
made to the scientific meetings of the Society as, on account of 
the nature of the plates required to illustrate them, are better 
adapted for publication in the quarto form. They are issued 
at irregular intervals. 

Fellows and Corresponding Members, upon payment of 
a Subscription of One Guinea before the day of the Anni- 
versary Meeting in each year, ai-e entitled to receive the 
Society's Publications for the year. They are likewise 
entitled to purchase the Publications of the Society at 25 per 
cent, less than the price charged for them to the Public. A 
further reduction of 25 per cent. is. made upon purchases of 
Publications issued prior to 1881, if they exceed the value of 
five pounds. 

Fellows also have the privilege of subscribing to the 
Annual Volume of the Zoological Hecord for a sum of 30i>. 
(which includes cost of delivery), payable on the 1st. of Julv 
in each year; but this privilege is forfeited nnless the 
subscription be paid before the 1st. of December following. 

The following is a complete list of the publications of the 
Society already issued. 



TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON. 

4to, 19 vols, and Index. 



'ol. 



;: < 

„ VII, 

„ VIII., 

„ IX., 

„ X., 

Index, Vols. 

Vol. XI., 
„ XII., 
„ XIII., 
„ XIV., 

;, XV., 

XVI 
„ XVII., 
„ XVIII. 
.. XIX. 



T., containing' r.O Plates. . . . (1833-35) 



71 
G3 

77 
07 
92 
73 
82 
99 
95 



I.-X 

containing 97 Plates 

;, 65 „ 

62 

1! 4/ ,, 

52 
„ 38 ,, 

41 
„ 43 „ 

24 



(18.35-41) 
(1842-49) . 
(1851-62) . 
(1862-66) . 
(1866-69) . 
(1869-72) . 
(1872-74) . 
(1876-77) . 
(1877-79) . 
(1833-79) . 
(1880-85) . 
(1886-90) . 
(1891-95) . 
(1896-98) . 
(1898-1901) 
(1901-1903) 
(1903-1906) 
(1907-1911) 
(1909-1910) 



Price to 

Fellows. 

£•3 13 6 



4 


. 




3 8 


3 




6 2 







5 4 


3 




11 6 







10 4 







9 8 


3 




12 1 


6 




10 


3 




7 


6 




9 12 







5 8 







6 8 


3 




5 5 







5 15 


6 




5 8 







5 18 


6 




4 1 







10 4 








Price to the 
Public. 

£4 18 Ot 

5 6 6t 

4 11 ot 
8 2 6t 

6 19 

15 
13 12 

12 11 

16 2 

13 7 
10 

12 16 

7 4 

8 11 
7 
7 14 
7 4 
7 18 

5 8 

13 12 



PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND 
CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF 



Price to 



LONDON. 8vo. 2 vols. (Letterpress only). ^^^^^^^^ 



Price to the 
Public. 



Part I. 
„ II. 



1830-31. 
1832. 



1 vol. 8vo 4s. Qd. 

4s. 6^. 



^t 



6s. 



PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON. 

8vo. 15 vols, (Letterpress only) and Index. (First Series.) 



Price to 

Fellows. 



Part I. 1833. 1 vol. 8vo. 4s. 6d. 



11. 1834. 

III. 1835. 

IV. 1836. 
V. 1837. 

VI. 1838. 

VII. 1839. 

VIII. 1840. 



4s. M. 
4s. 6d. 
4s. 6d. 
4s. 6d. 
4s. 6^. 
4s. 6d. 
4s. 6d. 



Price to the 
Public. 

.. 6s.t 
,, 6s. 



8vo. 



6s. 

6s, 

6s. 

6s.t 

6s.t 

13 vols, and Index 



Part IX, 1841 . 1 vol. 8vo. 4s. 6d 

X. 1842. „ 

„ XI. 1843. „ 

„ XII. 1844. „ 

„ XIII. 1845. „ 

„ XIV. 1846. „ 

„ XV. 1847. 
Index 1830-1847. 



Price to 


Price to the 


Fellows. 


Public. 


4s. 6d. 


., 65,t 


4s. 6^. 


,. 6s. 


4s. 6d. 


,. 6s.t 


4s. Gd. 


,, 6s, 


4s. 6d. 


.. Gs. 


4s. 6d. 


.. Bs.f 


4s. Gd. 


.. 6s.t 


As. Qd. 


., 6s. 



(Second Series.) 



Letterpress only. 
Price to Price to the 



With Plates coloured. 
Price to Price to the 



Part XVI. 1848. 

XVII. 1849. 

„ XVIII, 1850. 

., XIX. 1851. 

;, XX. 1852. 

„ XXI. 1853. 

XXII. 1854. 

„ XXIII. 1855. 

„ XXIV. 1856. 

XXV. 1857, 

„ XXVI, 1868. 

„ XXVII, 1859, 

„ XXVIII. 1860. 

Index 1848-1860, 



Fellows. 

1 vol. 8vo. 4s. 6^. 

„ 4s. 6f/. 

„ 4s. Gd. 
4s. Gd. 

„ 4s. Gd. 

„ 4s. Gd. 

„ 4s. Gd. 

„ 4s, Gd. 

,, 4s. Gd. 

„ 4s. Gd. 

„ 4s. 6^. 

„ 4s. 6^, 

„ 4s. Gd. 

,, 4s. Gd. 



Public. Fellows. 

Gs £1 8 

Gs 1 8 

6s 1 8 6 

6s 15 9 

6s 16 9 

6s 18 

6s 19 6 

Gs 1 8 6 

6s 1 8 

6s 1 8 

6s 1 11 6 

6s 1 11 6 

6s 1 11 6 

Gs. 



Public. 



. . £1 7 


6t 


..17 


6t 


. . 1 18 


Of 


..11 


Of 


..11 


Of 


..14 


Ot 


..16 


Of 


. , 1 18 


Ot 


..17 


6t 


..17 


at 


,.2 2 


Ot 


..2 2 


Of 


..2 2 


ot 



t Out of print. 
* In consequence of a re-arrangement of the stock of the ' Transactions,' the Society is 
now able to offer for sale, at the reduced price of £30, sets of Vols, v.-xvi. inclusive, and 
separate papere, of which a list can be supplied, at about one-fourth their published price. 



PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE 
ZOOLOGICAL SOCIETY OF LONDON. 8vo. 40 vols, and 4 Indices. 





Letterpress only. 


With Plates 


uncoloured. 


With Plates coloured. 




Price to 


Price to the 


Price to 


Price to the 


Price to 


Price to the 




Fellows. 


Public. 


Fellows. 


Public. 


Fellows. 


Public. 


18G1 


. 4s. Qd. . . 


. . Gs 


.. 95. 




.. 125 


. . 335. 


9d . 


.. 45s.t 


1862 


. 4s. Gd. . . 


.. Gs 


.. 9s. 




.. 125 


. . 335. 


9d. . 


.. 455. 


186:3 


. 4s. Gd. . . 


.. Gs 


.. 95. 




. . 125 


. . 33s. 


9d . 


.. 45s.t 


1864 


. 45. 6^. .. 


. . Gs.* 


.. 95. 




.. 125.t.... 


. . 335. 


9d. . 


. . 45s.* 


1865 


. 4 s. Gd. . . 


. . 6s.t . . . . 


.. 95. 




.. 125 


. . 335. 


9d. . 


. . 45s. 


1866 


. 4s. Gd. . . 


. . 6s.1. ... 


.. 95. 




.. 125.t.... 


. . 33s. 


9d. . 


.. 455. 


1867 






.. 95. 




.. 125.*.... 


. . 33s. 


9d. . 


.. 45s. 


1868 






.. 9s. 




.. 125 


. . 335. 


9d. . 


.. 45s. 


1869 






.. 9s. 




.. 12s 


. . 335. 


9d. . 


.. 45s. 


1870 






.. 95. 




.. 125 


. . 335. 


9^. . 


.. 45s. 


Index, 


1861-1870 




.. 45. 


6^/. . . 


.. 6s. 








1871 






. 9s. 




.. 125.*.... 
.. 125.*.... 


. . 335. 
. . 335. 


9d. . 
9d . 


. . 45s. 


1872 






.. 9i. 


.. 45s.t 


1873 






.. 95. 




.. 125 


. . 335. 


9d. . 


.. 45s. 


1874 






.. 95. 




.. 125.t.... 


. . 36s. 




.. 48s.t 


1875 






.. 95. 




.. 12s 


. . 36s. 




.. 48s. 


1876 






.. 9s. 




.. 125 

.. 12s 


. . 365. 
. . 36s. 




.. 485.t 
.. 48s. 


1877 






.. 9s. 


1878 






.. 9s. 




.. 125 


. . 365. 




. . 485. 


1879 






.. 9s. 




.. 12s 


.. 365. 




.. 48s. 


1880 






.. 95. 




.. 12s 


. . 365, 




.. 48s. 


Index, 


1871-1880 




.. 4s. 


Gd. .. 


. . 6s. 








1881 






.. 9s. 




.. 125 


. . 365. 




.. 48s. 


1882 






. . 9s. 




.. 125 

.. 125 


. . 365. 
.. 365. 




485 


1883 






.. 95. 


.. 48s. 


1884 






.. 9s. 




.. 12s 


. . 365. 




.. 48s. 


1885 






.. 95. 




.. 125 


.. 365. 




.. 485. 


1886 






.. Os. 




. . 12s 


. . 36s. 




.. 48s. 


1887 






.. 95. 




.. 12s 


. . 365. 




.. 48s.t 


1888 






.. 95. 




.. 12s 


. . 36s. 




.. 48s. 


1.889 






. . 95. 




.. 12s 

.. 125. .... 


. . 36s. 
. . 36s. 




. . 48s. 


1890 






.. 95. 


.. 48s. 


Index, 


1881-1890 




.. 45. 


6^. . 


.. 6s. 








1891 












. . 36s. 




. . 48s. 


1892 












. . 36s. 


. . 48s. 


1893 












.. 36s. 


. . 485. 


1894 












, , 36s. 


. -185. 


1895 












., 36s. 


. , 48s. 


1896 












. . 36s. 


. , 485. 


1897 












.. 36«. 


. . 485. 


1898 












.. 36s. 


. . . 48s. 


1899 












. , 36s. 


. . 48s. 


1900 












. 36s. 


. . 48s. 


Index, 


1891-1900 




. . 45. 


6f/. . . 


. . 8s. 






* No perfect cop 


les in stock. 








t Out of I 


)rint. 



PROCEEDINGS of the GENERAL MEETINGS for SCIENTIFIC 
BUSINESS OF THE ZOOLOGICxiL SOCIETY OP LONDON. 

8vo. 21 vols. 

Price to Price to the 

FellovFS. Public. 

1901, vol. 1 18s 24s. 

„ 11 18s 24s. 

1902, „ 1 18s 24s. 

„ II 18s 24s. 

1903, „ 1 18s 24s. 

„ li 18s 24s. 

1904, „ I 18s 24s. 

„ II 18s 24s. 

1905, „ 1 18s 24s. 

„ ' „ II 18s 2Js. 

1906, „ 1 18s 24s. 

„ „ II 18s 24s. 

1907, „ 1 18s 24s. 

„ ., II 18s 24s. 

1908, „ I 18s 24s. 

„ II 18s 24s. 

1909, ., 1 18s 24s. 

„ '„ n , 18s 24s. 

1910, „ 1 18s 24s. 

„ ., II 18s 24s. 

1911, „ 1 18s 24s. 



LISTS OF THE ANIMALS IN THE SOCIETY'S GARDENS. 

List of the Vertebrated Animals now or lately Living in the Gardens 

of the Zoological Society of London. (Eighth Edition.) 8vo. 

1883. Cloth, 4s. 6d. 

List of the "Vertebrated Animals now or lately Living in the Gardens 

of the Zoological Society of London. (Ninth Edition.) 8vo. 

1896. Cloth, 6s. 



Catalogue of the Library of the Zoological Society of London 
(Eifth Edition.) 8vo. 1902. Cloth, Os. 

THE OFFICIAL ILLUSTRATED GARDEN GUIDE— 9th Edition 
— with (1) a Railway and Street Map, showing a direct 
route to the "Zoo" from all parts of London and the Suburbs ; 
(2) a Plan of the Grounds, showing at a glance the location of 
the animals ; (3) a sbort description of some of the principal 
animals in the Collection (now coutaining over 3000 spe- 
cimens), together with 50 Photographic Illustrations and 
Index. Price Gc?. in Stiff Paper Cover, postage IgfZ., or in 
Green Cloth Cover price Is. 2d. post free. 



Regent's Park, London, N.W., 
June, 1911. 



P. CHALMERS MITCHELL, 

Secretary. 



These publications may he obtained at the Society's Office, 
at Messrs. Longmans'" {Paternoster How, E.G.). or through any 
bookseller. 



ox ABNORMAL ANTLERS OF A CARIBOU. 181 

EXHIBITIONS AND NOTICES. 

February 7, 1911. 

Prof. E. A. MiNciiiN, M.A., Vice-President, 
in the Chair. 

The Secretary read the following repoi-t on the additions made 
to the Society's Menagerie during the month of December, 
1910:— 

The registered additions to the Society's Menagerie during 
the montli of Dacember were 133 in number. Of these 76 were 
acquired by presentation, 27 by purchase, 12 were received on 
deposit, 17 in exchange, and 1 was born in the Gardens. 

The total number of departures during the month, by death 
and removals, was 216. 

Amongst the additions special attention may be called to the 
following : — 

1 Viverrine Cat (Felts viverrina), from India, 1 Northern Lynx 
(Felis lynx isabeUinus), from Tibet, 2 Binturongs (Arctictis hintu- 
rong), from Malacca, and 1 Yellow-throated Marten (Ifitstela 
Jlavigula), from India, purchased on Dec. 24th ; 4 Canadian 
Skunks (Mephitis mephitica) and 2 Little Skunks {Spilogale 
pittorius), fi'om IST. America, the latter new to the Collection, pur- 
chased on Dec. 29th. 

1 Silver Fox (Vulj^es argentatus), from Hudson's Bay, received 
in exchange on Dec. 12th. 

1 Feline Otter (Latra felina), new to the Collection, from La 
Plata, presented by William Clyne, Esq., on Dec. 6th. 

1 Brindled Gnu (Connochcetes tmtrimts), horn in the Menageiie 
on Dec. 1st. 

A collection of 15 Reptiles, including 1 Raddon's Lizard (Mahuia 
raddonii), 1 Smyth's Snake {GQ^ayia smythii), and 1 Yellow- 
throated Snake (Thrasops flavigidaris\ new to the Collection, 
from Dunkwa, Gold Coast, presented by Dr. H. G. F. Spurrell 
on Dec. 12th. _ 

Dr. C. W. Andrews, F.R.S., F.Z.S., exhibited the skull of a 
Sabre-toothed Tiger (Smilodon californicus), from an asphalt 
deposit in California, and pointed to anatomical characters which 
tended to prove that the animal used its large canines for stabbing 
and tearing, not for biting. 

The Secretary exhibited a mounted specimen of the Platypus 
( Orniihorhynchus jjaradoxus\ which had been lent for the purpose 
by Mr. P. St. Michael Podmore, F.Z.S. 

Mr. Edward Gerrard, F.Z.S., exhibited the head of a Caribou 
(Rangifer tarandns caribou) (text-fig. 52), shot by Sir John Rogers, 
K.C.M.G., in British Columbia, which had a distinct forked third 
antler over the centre of the orbital arch of the frontal bone just 

Proc. Zool. Soc— 1911, Ko. XIII. V6 



182 



ox AliXOUJIAL AXTLE1?,S OV A C^AIUBOU. 



in front of the normal antler, which carried a single anterior basal 
tine, and separated from it by a strip of hairy skin. There was a 
slight burr on a, level with the skin, but no pedicle. It had the 
appearance as if it could be shed in the same manner as the 
normal antlers. 

Text-fiiT. 52. 




Head of a Caribou (Banr/ifer tarandus caribou) with abnormal antlers. 

A very fine head of an Eland {Taurotragus derhianus gigas) 
obtained by Major Gordon on the Bahr-el-Ghazel, and three fine 
heads of White-tailed Deer (Borcelaphus aniericanas macrurics) 
Avere also exhibited, one pair of antlers being abnoiinal. 



PZ S 1911 Pill 




V,(}' H^> ^7x500 ^% 8^500 



WN.F.W. del. 



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GAS GLANDS OF TELEOSTEI. 



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.Wilson, Cambridge . 



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GAS GLANDS OF TELEOSTEI 



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GAS GLANDS OF TELEOSTEI. E.WHson^Carnbridge. 



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GAS GLANDS OF TELEOSTEI. 



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GAS GLiVNDS OF TELEOSTEI. 



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GAS GLANDS OF TELEOSTEI. 



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GAS GLANDS OF TELEOSTEI. 




E. Wilson, Gaml)ridge . 



ox THE GAS GLANDS OF SOME TELEOSTEAN FISHES. 183 



PAPERS, 

11. On the Structure and Function of the Gas Glands and 
Retia Mirabilia associated with the Gas Bkdder of some 
Teleostean Fishes, with Notes on the Teleost Pancreas. 
By W. N. F. Woodland, F.Z.S., The Zoological 
Department, University College *, London. 

[Received November 9, 1910 : Read February 7, 1911.] 
(Plates Il.-IX.t and Text-figures 53-62.) 

OONTE^TTS. 

Page 

Introductory 183 

Tart I. — The Structure of ''Red Bodies " 184 

Angiiilla as a Type 185 

Other Examples of this First Type of "Red Body" — Myriis, Opldchthys. 191 

A Second Type of "Red Body," exemplilied by Syngnathus 193 

Other Examples of this Second Type — Gohiits, Siphonostoma, Nerophis, 

GasterosieuSs Peristethiis, Tvigla, Smarts 193 

Xotes on the Teleost Pancreas 198 

A Third Type of " Red Body," exemplified by Ojakidium SOi 

Other Examples of this Third Tj-pe — Bo.v, AtJieritta, Coris, Covvum, 

Sargus, Balistes, Caranx, Serraiius, Zens, Gadus, Cepola, Perca . 205 

The Classification of Gas Glands 219 

The Giant Cells of Gas Glands 210 

The Gas Bubbles iu Gas Gland Cells ., 198 

Summary of Part I. and General Conclusions respecting Gas Gland 

Structure 215 

Part II. — The Physiology of "Red Bodies" 220 

The Principal Theories as to the Origin of the Bladder Gases 220 

Thilo 221 

Moreau 222 

J aeger 223 

Nusbaum and Reis 223 

Discussion of the Views of Jaeger and Nusbaum & Reis 22i 

A Statement of the Modern View of the Physiology of the " Red 

Bodies " 228 

Some additional Suggestions by the present Writer — the Rationale of 
the Rete Mirabile, the Disintegration of the Red Blood Corpuscles 

and other Minor Features of "' Red Body " Structure 230 

Appendix A. — The Sources and Modes of Preparation of Material 236 

Appe>-dix B. — On the Artificial Production of Gas Bubbles in Cells of the 

Gas Gland 237 

A ddenda 241 

Literature Lists 242 

Introductory. 

In the present paper it is proposed to give a fairly complete 
account of the gross and histological structui-e of the vasculo- 
glandular organs or '' red bodies " found in the wall of the gas 
bladder t of many teleost fishes, and to ofier certain suggestions 

* The greater part of the work t-onnected with the present paper was carried out 
in the Zoological Department of King's College, London. 

t For explanation of the I'lates see pp. 245-248. 

X The terms " swim-bladder " and " air-bladder " are inappropriate, since fishes do 
not use this apparatus for swimming and it does not contain air. 

13* 



184 DR. W. X. F. WOOELANB ON THE 

respecting the physiology of these organs. Though the litera- 
ture * concerned with the morphology and physiology of the gas 
bladder is, of course, very considerable, yet up to the present only 
one author (6i, 65) has attempted a complete review of the 
subject of the present paper in the light of recent work. Further, 
although there exist several memoirs dealing with special types of 
" red body," yet most of these are so occupied with cytological 
and other details that, in the opinion of the present wiiter, they 
have failed to supply such an account and such illustrations of 
the gross structure of these " red bodies " as to render evident 
some of their most remarkable features. Finally, the Jaeger- 
Nusbaum controversy calls for an independent description of these 
unique organs, so interesting from the physiological standpoint. 

I have divided the present paper into two parts, the first of 
which deals with the structure and the second with the physio- 
logy of the " red bodies." With reference to the drawings 
illustrating Part I., I have, with a few exceptions, adopted a 
definite scheme of coloration in order to facilitate comprehension. 
Green has been adopted to denote the internal lining epitiielium 
of the bladder, and therefore the glandular, i. e. gas gland, epi- 
thelium is always of this colour in the drawings, since this is but 
a special development of the usually squamous lining epithelium ; 
red denotes blood ; black or grey has been employed for connec- 
tive and muscular tissue and secretion mateiial ; and yellow has 
been used to indicate the pancreas, which is often closely 
associa^ted in a mechanical sense with some parts of the " red 
body." The sources and methods of preparation of my material 
are stated in Appendix A (p. 236). 

Part I. — The Structure of the "Red Bodies" in 

VARIOUS TeLEOSTEI. 

It is necessary to recognize at the outset that the term " red 
body " unfortunately has been applied to difi'erent structures by 
no means equivalent to each other : thus, e. g., the " red bodies " of 
the Common Eel are simply the two retia mirahilia, whereas the 
"red body" of the Cod consists both of numerous small retia 
mirabilia and of the large mass of epithelial cells called the gas 
gland, the retia and the gas gland being, in this case, intimately 
a,ssociated. A gas gland exists in the Eel as in all other fish 
with " red bodies," but it is not in close connection with the 
rete mirabile as in the Cod. This being the case, it must be 
understood that by the term " red body " all that is meant is a 
red mass situated in the bladder-wall, the constitution of which 
is variable. I may add that in the vast majority of cases " led 
body " includes both i-ete mirabile and gas gland. As an intro- 
duction to the study of gas gland structure I shall first describe 
the anatomy of one of the most simple as well as most common 

* For the lilevatuveand historical reviews of the entire svrhject of gas hladders up 
to 1806 see Miliie-Edwarcls (49) and Gonriet (34) ; for more recent literature sec 
the lists supplied by Bridge (22), Hiifner (38), Jaeger (44), Deineka (29), Keis & 
Is'usbauui (62, 63), and at the end of the present pai)er. 



GAS GLANDS OF SOMK TELEOSTEAN FISHES. 185 

forms, viz. tliat of the Common Eel, Angailla valyarls. The " red 
body " of the Eel has been described several times previously, 
e. g. by Jacobs (42, 43), Quekett (61), and Corning (38), but never, 
I ve]iture to think, in a sufficiently clear or comprehensive 
manner and certainly never adequately illusti'ated. 

The Vascwlo-glcmdalar Structure hi the Bladder of the Common Eel 
(Anguilla vulgaris). 

It is well known that the bladder of the Eel is a.n elongated 
sac lying dorsally and towards the posterior end of the body 
cavity and connected with and opening into the oesophagus by 
means of a long wide duct — the ductus pneumaticus (text-fig. 53, 
P.D., p. 186). The pneumatic duct opens into the bladder at about 
midway in its length, the bladder in consequence being divisible 
into anterior (A.B.) and posterior (P.B.) regions. The bladder 
and duct are both situated in the same horizontal plane, ?'. e. the 
duct opens into the bladder at the side, not dorsally or ventrally. 
Running along the vential side of the duct are an artery and 
a vein situated close together (A.Y.). This artery and vein give 
rise to the two ovoid retia mirabilia (R.M.) where the duct opens 
into the bladder, the two retia lying one on each side of the 
duct and therefore being dorsal and ventral in position respec- 
tively and in the same vertical line. 

The wall of the bladder (see PI. II. figs. 1-6, e.g.) in fish generally 
is best described as consisting of the tunica externa and tunica 
interna. The former consists of two definite layers of fibrous tissue, 
the outer being loose in texture, often containing muscular tissue 
and varying greatly in thickness, the inner dense, silvery in appear- 
ance, aiid principally consisting of elastic fibres. The latter consists 
of a thin basis of dense connective tissue, with wluch is closely 
connected the innermost layer of the bladder wall, a layer con- 
sisting usually of flattened cells. It is this innermost cellular layer 
which gives rise to the mass of glandular cells in connection 
with a rete mirabile — the gas gland. The tunica interna is 
easily separable from the tunica externa, and it is advisable to 
remove this latter before cutting sections of the gland. 

The vascular supplj' of the bladder may first be considered. It 
has already been mentioned that an artery and a vein run along 
the venti'al side of the wall of the ductus pneumaticus. These 
two vessels are situated in the outer fibrous layer of the tunica 
externa (PI. II. fig. 1), which is here of considerable thickness, and 
they supply several arteries to the duct and receive several veins 
from it. Shortly liefore these two vessels reach the region of the 
jimction of the duct with the bladder, the artery and the vein 
each divide into two (fig. 2), so that there are two pairs of vessels 
instead of one, each pair consisting of an arteiy apd a vein. 
ISTow the artery and vein of each of the.se pairs undergoes further 
subdivision, the smaller arteries and veins resulting fi'oni this 
subdivision intermingling in their course, but not once' com- 
municating \\\\X\ each other, so that we now have situated towards 



186 



DK. W, W. F, WOODLAND ON THIS 



the upper and lower sides of the ductus pueumaticus two distinct 
masses of intermingled arteries and veins (fig, 3), each mass 
having resulted from the subdivision of the two pairs of arteries 
and reins, which, as we have seen, have themselves originated 

Text-fiff. 53. 



A- 



p.a- 



R.nr 



P.B: 



A.B. 



The bladder, duct, and "red bodies " of Anguilla vulgaris, 
viewed from the side (after Quekett). 

P.D., pneumatic duct which opens anteriorly into the a?sophagus; A and V, artery 
and vein supplying bladder; E.M., rete mirabilej A.B., anterior region of 
bladder; P.B., posterior region of bladder. 



GAS GLAXDS OF SOME TELEOSTEAX FTSllES. 



187 



by tlie division of the initial artery and vein. The subdivision 
of the arteries and veins in each mass proceeds rapidly, until 
there is forined by this process ari ovoid mass of extremely fine 
parallel arterial and venous capillaries, closely intermingled Avith 
each other, but never intercommunicating and indistinguishable 
structurally as arteries and veins under the highest powers of 
the microscope (fig. 4). These two ovoid masses of parallel 
arterial and venous capillaries lying on the dorsal and ventral 
sides of the ductus pneumaticus just a.nterior to its junction 
with the bladder are the retia mirabilia of the eel-bladder (text- 
fig. 53, R.M.). 

The finest arterial and venous capillaries of each rete mirabile 
run parallel to each other for a short distance, but soon they 
commence to unite, arterial capillaries with arterial capillaries 
and venous with venous, in order to re-form larger vessels similar 
to those which pi-oduced the capillaries by subdivision. This 
process of reunion proceeds so far as to form two masses of 
lai'ge intermingled arteries and veins similar to those depicted 

Text-fig. 54. 




External aspect of a rete mirabile (after Quekett). The anastomoses of the 
vessels shown in this figure do not occur in actiiality. 

in figure 3 (c/. figs. 3 & 5). When reunion has proceeded thus 
far, each of these masses of re-foi-med arteries and veins once more 
breaks up into fine capillaries in connection with the vascular 
supply of the glandular folded epithelium which lines the bladder 
internally. I reproduce here as faithfully as I am able (text- 
fig. 54) Quekett's figure (61) of the construction of the Eel's rete 
mirabile, which illustrates in a general way that which I have 
just described. Text-fig. 54 is vm satisfactory in that it does not 
indicate the fact that the vessels and capillaries forming the 
rete are of two kinds — arterial and venous — closely intermingled 
with each other, although never intercommunicating ; also anas- 
tomoses never occur between adjacent large vessels in the manner 
shown in the figure. A more satisfactory diagram of the 
construction of the Eel's rete mii'abile is the one I have 



188 



DR. W. N. F. WOODLAND ON THE 



designed for text-fig. 55. In this the axtery and the vein 
giving rise to a rete mirabile are distinguished from each other 
by the latter being cross-striped, the subdivision of each of 
them is indicated, also the intermingling of the fine parallel 
capillaries, their reunion and the second breaking up into capil- 
laries of the large arteries and veins thras formed to supply the 
epithelium of the gas gland. 

Up to the present I have spoken of the arterial and venous 
capillaries or vessels composing each of the two retia mirabilia 
merely as tubes, but if, of course, we take into account the 
direction of the flow of blood in the two sets of vessels, then 
it is evidently incorrect to speak of the ai-teries and veins both 
subdividing at the anterior pole of the rete mirabile to prodwce 

Test-fiff. 55, 




Cr.E. 

The constrmction of tile rete mirabile bipolare geminum (R.M.). 
G.E., glandulai epithelium of gas gland. 

capillaries : what is really happening in this region is that the 
?<rteries are subdividing and the veins uniting (text fig. 55). 
That this is the actual mode of construction of the rete mil a bile 
is not only proved by the study of serial sections but also by the 
injection method. And here let me I'emark upon the extra- 
ordinary process implied by this consti'uction of the rete 
mirabile — a term which, like its German rendering, "wvmder- 
netze," happens to be peculiarly appropriate*. It is implied 
that parallel vessels carrying blood in opposite directions can so 
influence each other, or be so influenced by a common cause, as 
to break up into a rete mirabile at the same spot, the subdivision 
of each of the two sets of vessels being so adapted to the other 
as to ensure the close apposition and intimate intermingling 
of their finest capillaries f. It is evident that this intricate 

* Miiller (53) termed tins type of rete mii-ahile a " rete mirahile bipolare gemiimm," 
in contradistinction to a " rete mirabile bipolare simplex," such as the purely arterial 
carotid srland of Amphibia. 

t The fact illustrated bj' these retia that blood-vessels are able to capillarize on 
their own account, i. e. quite apart from any other tissue, first attracted my attention 
to these " red bodies " when engnsred in certain speculations concerning the "renal- 
jiortal system " in Vertebrates (76, 77). The formation of the retia seems to lend 
support to Dr. Shore's views (68, 69) concerning the vascularization of the liver 
as distinguished from the venous penetration of the kidney-substance to form the 
"renal-portal system." I hope shortly to investigate the subjects of the"renail- 
■portal system'' and the various kinds of retia mirabilia from a practical standpoint. 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 189 

mechanii^m must he for some definite purpose, especially since it 
is found in all fishes possessing gas glands. What that purpose 
may be is a question I shall discuss later (see Part II.). 

The epithelium lining the ductus pneumaticus, Avhich is sup- 
plied with a few branches direct from the bladder artery and is 
therefore not supplied by the vessels connected with the retia 
mirabilia, is quite simple in character. It merely consists of a 
single layer of flattened cells raised up at intervals by the large 
capillai'ies running in the basis of connective tissue underlying 
the epithelium (PL II. fig. 7). These capillaries in the wall of the 
duct apparently have much the same surface ari-angement as those 
found in the lungs of certain reptiles (Quekett, 61), but it is 
hard to suppose from this fact alone that the epithelium of the 
duct is normall}^ respiratory in function, since the blood supplied 
to this has already been oxygenated in the gills ; on the other 
hand, it is equally improbable that the ai'terial blood in the wall 
of the duct serves to alter the composition of the adjacent gas 
since there is reason to suppose that the duct normally serves as 
a mere exit for the superfluous gas of the bladder (Jaeger, 45). 
It is possible, however, that this large vascular supply of the duct 
epithelium in the Eel is really correlated with the terrestrial 
habits of the animal, since when an Eel travels across a meadow 
the gills are useless for obtaining oxygen, and under these circum- 
stances the animal doubtless draws upon the oxygen in the 
bladder — the duct acts as a lung, Moreau proved that Perch, 
when placed in water previously boiled so as to deprive it of 
dissolved air, utilized the bladder- oxygen for purposes of respira- 
tion. An interesting confirmation of this view is that in other 
physostomes {Myrus, Ojyhichthys, Esox) the dtict epithelium is, as 
we shall see, practically devoid of capillaries *. 

The lining epithelium of the bladder proper is very difierent 
in chai'acter from that of the duct. It consists of a more or less 
folded single layer of cells which are large in size and short 
columnar in form, slightly gTanular, and, when stained by the 
picro-indigo-carmine method (Appendix A), usually rather opaque 
(PL II. fig. 8). The folding of this layer is due to the presence of 
large capillaries which push their way inwards towards the bladder- 
cavity from the connectiA^e tissue base, and in so doing cause the epi- 
thelial layer to form corresponding emergences. Thus the iutei'ior 
of each projection of the glandular epithelium into the bladder 
cavity is occupied by an extension of connective tissue enclosing 
capillaries. The spaces situated between such emergences adjacent 
to each other are also regarded as forming gland ducts. The 
folding of the epithelium is very vaiiable in difierent parts of the 
bladder wall, in some parts almost disappearing and in others 
being more accentuated than that depicted in figure 8 ; this 
folding is much more marked in the anterior region of the 
bladder than in the posterior. It might appear to be significant 
that immediately the lining epithelium of the duct becomes sup- 
plied with blood connected with the rete mirabile system of vessels 

* I iiiul that Jacobs (42, 43) status tliis view as a fact. 



190 DR.. W. N. F. WOODLAND ON THE 

(that is, M*laen the duct opens into the bladder) it then assumes 
the glandular folded character of the bladder epithelium, were it 
not for the fact that in other physostomous genera this sharp 
distinction between duct- and bladder-epithelium is absent, as I 
sha,ll show shortly. 

The intracellular structures described in detail below as being 
present in the cells of the gas glands of other teleost fishes — 
intracellular gas bubbles, capillaries, and ducts — are not easily to 
be detected in my preparations of the Eel, but I believe I have 
seen ga,s bubbles present in a few cases, also the capillaries 
occasionally intrude somewhat upon the cells at their bases. 
Granular matter, ejected into the bladder- cavity with the gas, is 
so small in aanount in the gland ducts of my preparations that it 
may be said to be absent. 

The folded glandular epithelium of the gas gland of the Eel, 
and presumably of all other genera possessing a folded epithelium, 
originates from a single unfolded layer which during development 
becomes pushed out into the folds by the activity of the large 
ca.pillaries situated in the thin layer of connective tissue- lying 
at the base of the epithelium *. 

Thus in the type of " red body " found in the Common Eel 
we have two large retia rairabilia,, situated some distance from 
the epithelium of the bladder which they supply, and the greater 
part of this lining epithelium of the bladder is glandular and 
folded in a simple manner to form the gas gland. In all " red 
bodies " we have these two elements, the rete mirabile and the gas 
gland, but in the majority of " red bodies " these two elements 
are much more intimately associated with each other and more 
complex in form as compared with the structure of the simple type 
of " red body " just described. The Eel type of " red body " serves 
as a convenient starting-point for considering more complex types. 

Before considering these types, it is necessary to point out a 
serious mistake contained in almost the only English paper 
dealing with the present subject, that of Vincent and Barnes (75), 
which, though the paper is now quite out of date, has found its 
way into at least one recent text-book (' Cambridge Natural 
History,' vol. vii. 1904, p. 308). Vincent and Barnes, following 
Coggi (25), state that " red glands are only found in those 
bladders which are devoid of a ductus pneumaticus," and they 
draw a sharp distinction between Physoclisti, which possess the 
gas gland, and Physostomi, which are supposed to be devoid of 
one. I need hardly say that no such distinction exists. Vincent 
and Barnes are right, in so far as they say that the " red bodies " 
of the Eel are only partially comparable with those of the 
Gadidse {vide supra), but it is difficult to understand how they 
came to overlook the large and conspicuous glandular epithelium 
lining the bladder cavity. Further, as Deineka (29) points out, in 
the physostomous Pike {^Esox hocitos) a conspicuous gas gland 

* I am much indebted to Dr. W. G. Ridewood for kindly presenting mc with 
most of my young material. 



GAS QLANJDS OF SOME TELEOSTEAN FISHES. 191 

exists (described by Coggi, 26), compaxable in all essential respects 
with that of the Cod. Also Vincent and Barnes, unlike many of 
their predecessors, altogether fail to appreciate the peculiarity of 
the arterial and venous nature of the rete mirabile ; indeed, they 
even appear to have some doubt as to the fact itself. They say : 
'' Corning has discriminated betwee'U arteries and veins in what 
Ave have usually spoken of as the ' capillary masses.' This 
he has done by means of a, sei'ies of injections. .... This method 
would certainly give some idea of the distinction between the 
smallest arterioles and smallest veTiules, but we are doubtful 
whether it would be more than a. very rough method of distinc- 
tion, that it would be, in any given case, an infallible test as to 
whether a particular small vessel were to be called artery or vein. 
Much would depend, it appears to us, on the strength of the 
injecting force [!J. However this may be, we have failed to make 
out anything in the minute structure which would warrant us in 
dividing up the ' capillary masses ' into arterioles and venules." 
This statement suggests that Vincent and Barnes never worked 
through a series of sections of the Eel (or any other similar) 
" red body " (which possibly also accounts for their missing the 
gla,ndular epithelium of the Eel, which, as above stated, lies 
separate from and posterior to the retia mirabilia *), since the 
artei'ial and venous nature of the rete mirabilia is rendered quite 
evident by this means alone ; also the last-quoted statement of 
Vincent and Barnes is still less comprehensible when we remember 
that these " red body " retia mirabilia have been correctly 
described in full and compared with the several other kinds 
of retia mirabilia which exist by Johannes Mliller (53) in 1840, 
not to mention Owen's description (not confirmed by his figure, 
however) of the " i-ed body " of the Eel just referred to f. 

(/ Vasculo-glandular Structures of the Anguilla or First Type in the 
Bladders of some other Anguilliformes. 

Myrus vulgaris Kaup. 

In this physostome the vascular and glandular conditions are 
similar to those found in Anguilla vtdgaris. The only essential 
distinction between the two is in the character of the lining 
epithelia of the duct and bladder, and in this particular the 

* A fact dearly stated hy Owen (' The Comparative Anatomy and Physiology of 
Vertebrates,' vol. i. 1866, p. 495) : " The two chief ' retia mirabilia ' or vaso- 
ganglions, in the air-bladder of the Eel and Conger, which are situated at the sides 
of the opening of the air-duct, are also ' bipolar ' and consist of both arterioles and 
venules [curiouslj' enough, however, in contradiction of this statement, the figure 
329 of the ' vaso-ganglion ' of the Eel supplied on p. 496 shows the blood going 
through all capillaries in one direction !] : they consist of straight parallel capillaries, 
as in fig. 329 ; their afferent trunks do not ramify in the immediate margin of the 
vaso-sanglion from which they issue, as in the vaso-ganglions of the Cod, Burbot, 
Acerine, and Perch, but run for some distance before they again branch to form the 
common capillary system of the lining membrane of the air-bladder." 

t These criticisms must not be held to imply any lack of appreciation of a very 
lucid and interesting paper ; the fact, however, that the statements criticised have 
been made use of by the late Professor Eridge so recently as 1904 shows the need for 
correction. 



1 92 DR. W. N. F, WOODLAND ON THE 

distinction is striking. In Aiiguilla the epithelia of the duct and 
bladder are, as we have seen, quite different in character; in 
Myrus vulgaris, on the other hand, they are very similar. The 
duct epithelium consists of a single layer of short columnar cells 
which is only slightly folded. Yery few capillaries are present in 
the connective tissue layer underlying the epithelium, though 
here and there large capillaries are to be seen in transverse 
section. Thus the duct is not lined by squamous epithelium as 
in Anguilla. In the bladder of Myrus vulgaris tlie epithelium is, 
as just stated, similar to that of the duct, the only difference 
being that the cells are larger. The folding of the epithelium is 
at most very slight and most pronounced in the region of the 
opening of the duct ; in the anterior arid posterior regions of 
the bladder, folding of the epithelium is practically absent. Tliere 
is never any approach to the degree of folding seen in Anguilla. 
It is also noticeable that where the epithelium passes over a large 
capillary (and capillaries are not very numerous) the cells 
immediately assume a more squamous character. 

Ophichthys (Sph^gebranchus) imberbis Delar. 

In this physostome the plan of construction of the " red 
body " is essentially the same as that found in Anguilla and Myrus, 
but there exists in the single specimen at my disposal one 
modification in connection with the rete miiabile. When 
the bladder is viewed in situ, a single "red body" is seen to 
be present which is situated anteriorly in the ventral wall where 
the duct joins the bladder. A series of transverse sections shows 
that this "red body'' is, as in the Eel, a large rete mirabile, 
which, however, is disposed in a different manner in the present 
instance. The artery and vein run along the right side of the 
bladder and break up to form the rete in such a manner 
that the long axis of this lies transversely to the length of 
the fish (PI. II. fig. 9). The capillaries of the rete mirabile 
unite as usual to form larger vessels, which here, of course, lie to 
the left side of the bladder, where they break up anew to supply 
the epithelium of the gas gland. As in Myriis, there exists no 
sharp distinction between the glandular epithelium of the duct 
and that of the bladder, also the capillaries in connection with 
the epithelia are comparatively few in number. This epithelium 
consists, as in the two preceding genera, of a single layer of 
moi'e or less columnar cells which occasionally becomes A'ery 
slightly folded, but over the greater pai-t of the wall, both of the 
bladder and the duct, the epithelium is a simple unfolded layer of 
columnar glandular cells. Strange to say, the cells of the epi- 
thelium of the bladder in my preparations are distinctly smaller 
than those ol the duct. Occasionally large capillaries are present 
in the bladder wall, and where these occur the epithelial cells 
become, as in Myrus, almost squamous in shape. Small capillaries 
also exist, though they are not numerous. The epithelium of the 
duct remote from the bladder becomes much more squamous in 
character than elsewhere. 



gas glands of some telbostean fishes. 193 

Ophichthys (Ophisurus) serpens L. 

The general plan of the tioo retia mirabilia and the gas gland 
is exactly that found in Angailla. The epithelial lining is, as 
ill Angailla, squamous and covers projections of the subjacent 
connective tissue, which contain, not one large capillary, but many 
small ones. Posteriorly the lumen of the duct, which anteriorly 
is circular in transverse section, becomes very I'estricted and star- 
shaped in outline, owing to the great size of the connective tissue 
projections just mentioned — a feature not met with in the species 
described above. The epithelium of the bladder wall is hardly 
folded at all over the greater part of its area, simply consisting 
of a single layer of short columnar glandular cells, but in places 
folding occurs, occasionally to such an extent as to resemble 
the bladder epithelium of Angicilla, but this is rare. 

It is curious how the epithelial linings of the duct and bladder 
differ in these four closely related fish — Aiiguilla, Myrus, Ophichthys 
imberbis, and 0. serpens. In the duct and bladder of Anguilla Ave 
have the extremes of flattened and folded epithelia ; in the other 
genera intermediate conditions of the epithelium exist, both ns 
regards the form of the cells and their arrangement, and the duct 
and bladder epithelia are not sharply distinguishable from each 
other, and yet the vascular supply has the same distribution in all 
cases, although it undoubtedly varies considerably in amount, 
being much greater in the Eel than in the other genera. The 
diflTerent conditions of the rete mirabile in the two species of 
Ophicthys are also noticeable. 

The ty^Q of vasculo glandular apparatus found in the bladders 
of the preceding genera may be defined as follows : the glandular 
epithelium is composed of a single layer of cells which either 
remains unfolded or is only simply folded, and the retia mirabilia 
a,re one or two in number and are large and distinctly sepai-ate 
from the gas gland, the capillaries of the rete reuniting to form 
large arteries and veins before coming into connection with the 
glandular epithelium. 

A Second Type of Bladder Vasculo-glandidar Apparatus, 
as exemplified in Syngnathus acus Mich. 

Syngnathus acus is a physoclistous fish in possession of a " red 
body," which is, on the whole, very similar in construction to that 
of the Eel, but which nevertheless exhibits certain differences 
which justify us in separating it as a distinct type. The "red 
body " in Syngnathus is visible with tbe naked eye as a small 
red patch at the extreme anterior end of the bladder. An artery 
and vein running in the anterior attachment of the bladder divide 
up in the manner already described for the Eel to form the single 
rete mirabile (Pis. II. & III. figs. 1 0-14). It will also be observed in 
figures 10 and 11 that posterior extensions of the diffuse pancreas 
(indicated in the figures by yellow masses) penetrate into the 
anterior end of the I'ete mirabile and, as will be shown in detail 



104 DR. W. X, F. WOODLAXD OX THE 

later, are closely associated with the larger veins. Lying along- 
side the rete mirabile is the anterior extremity of the elongated 
closed bladder (fig. 14), the wall of the anterior region of which 
is very glandular and vascular. A little way behind the anterior 
exti-emity of the bladder, the side of the rete next the bladder 
comes into connection with the bladder epithelium (fig. 15), the 
capillaries uniting (i. e. the arterial capillaries are uniting and 
the venous capillaries are really here subdividing to form the 
minute venous capillaries of the rete) to form slightly larger 
vessels before supplying the epithelium of the gas gland. This 
process of supplying the glandular epithelium goes on until the 
whole of the rete mirabile is used up (PI. III. figs. 15-17,), the 
fine capillaries of the rete always uniting to some extent to form 
larger vessels before these supply the epithelium. The bladder 
epithelium, as shown in the figures, becomes enormously thickened 
by a process of folding during the breaking-up of the rete mira- 
bile. Posterior to the rete mirabile, which has thus disappeared 
in supplying the glandular epithelium, this latter itself gradually 
dwindles (fig. 18) until in the posterior region of the bladder 
the cavity is lined with simple squamous non-glandular cells. 

The epithelium of the bladder is quite simple in type and 
essentially resembles the folded epithelium of the Eel, though 
the folding is much more pronounced. Figure 19 represents the 
glandular epithelium in the unthickened region, that is, lining 
any side of the bladder cavity other than that adjacent to the 
rete mirabile, where, as just stated, the epithelium is much 
thickened. The folds of the epithelium due to the large capil- 
laries which push out the epithelium are so deep that we have 
here, as in Anguilla, distinct ducts fonxied between the folds 
which contain on occasion large quantities of floccular matter 
evidently derived from the cells. The thickened epithelium 
adjacent to the rete mirabile is formed by the great ex- 
tension of the folds just mentioned, the cells of which often 
join at intervals, i. e. the folds run together so that the at-first- 
separate ducts form anastomosing channels before opening into 
the bladder. The cells forming this epithelium are faintly 
granular and almost cubical in form, and here and there contain 
gas bubbles (not shown in the figure) and intracellular capil- 
laries, but since I have examined these structures in greater 
detail in other types, I shall not refer to them any further at 
present. I shall also, and for the same reason, merely mention 
here the conspicuous opacity of the cytoplasm of the gland cells 
which comes into immediate contact with a capillary (fig. 19). 

One other fact which is very well shown in my slides of 82/n- 
gnathus remains to be mentioned, and that is the disintegration 
of the red blood corpuscles contained in the cajoillaries of the 
rete mirabile and the gas gland first into spherical globules and 
ultimately into granular matter. This granular matter resulting 
from erythrocytolysis is, of course, only found in the blood-stream 
and is to be carefully distinguished from the granular matter 
above described as being found in the bladder lumen and gland 



GAS GL.VNUS OF SOME TKLKOSTEAX FISHES. 195 

ducfcs just external to the cells of the gii,s gland. Figure 21 
(PI. III.) shows several stages of this disintegration of the 
erythrocytes, the significance of which I shall discuss later. 

Figure 20 represents a highly-magnified transverse section 
through the fine arterial and venous capillaries composing the 
mass of the i-ete mirabile. The two kinds of capillaries are 
quite indistinguishable by mere inspection. The capillaries are 
separated from each other by a small amount of connective tissue. 
In the very young fish, l. e. before the capillaries have subdivided 
to the extent found in the adult, the ai-terial ca.pillai'ies are dis- 
tinguishable from the venous, the former being thicker- walled and 
somewhat smaller in calibre (see also Bykowski & Nusbaum, 23). 

Thus the chief features of the second or tSyiignathiis type of 
"red body" are that the glandular epithelium is arranged in 
simple folds, that it is restricted in aren, not lining the whole of 
the bladder cavity, and that the rete mirabile is contiguous with 
the gas gland, although a small amount of reunion of the capillaries 
of the rete may occur before these supply the epithelium. 

Other ExMiivples of the Syngnathus Ttjjje of''' Red BodyT 

GOBIUS NIGER. 

As another example of the second or Syngnathus type of 
bladder vasculo -glandular apparatus I shall now describe that 
of Gobiits niger caught at Naples*. The " red body " of Gohius 
is a small oval red patch situated just anterior to the centre of the 
oval outline of the bladder (PI. III. fig. 22). It consists of a single 
rete mirabile which supplies a small area of glandular ej^ithelium. 
As seen in a series of transverse sections, the artery and vein, 
Avhich break up to form the rete, at first lie outside the several 
layers of the bladder wall (PI. IV. fig. 23) together with large 
masses of pancreas. The artery and vein subdivide as usual to 
form a cluster of small arteries and veins, which in Gohnis, as in 
Syngnathus, are intermingled with masses of panci-ea-s (fig. 24). 
It will also be noticed that this cluster of arteries and veins is at 
this stage in process of breaking through the thin dense outermost 
layer of the bladder toall, so as to lie within this latter, large 
masses of pancreas being included with the cluster of vessels. 
Still moi-e posteriorly when the arteries and veins have undergone 
subdivision to a greater extent, the whole mass of vessels together 
with the included portions of pancreas have become almost 
entirely included loithin the bladder wall, small portions of pan- 
creas only being left outside (fig. 25). Finally, when the rete 
mirabile is completely formed it is wholly included within the sub- 
stance of the bladder wall, as in previous genera (fig. 26). The 
rate mirabile supplies the adjacent area of glandular epithelium 

* Grohiiis paganellus, obtained from Plymouth, possesses a star-shaped " red bodv " 
resembling tiiat of Perlstethus in constructiou (PI. V. fig. 37, and text-tig. 57), 
though less definite in outline. Gohius minutus, also obtained from Plymouth, has 
a, similar star-shaped " red bodj'," but the gas gland epithelium in some specimens 
is hai-dh' folded at all, merely consisting of a single layer of cells oncloaing- large 
capillaries ; in others it is folded as in Lrobitts iiiijcr (PI. IV. fig. 27). 



196 DI'o W. N. F. WOODLAND ON THE 

ill the usual way, the epithelium lining the rest of the bladder 
remaining squamous in character. It is, however, noticeable 
that in the rete of Gobius the fine capillaries do not unite 
too'ether at all to form larger vessels before supplying the 
o-landular epithelium (see fig. 27). In Syngnathus it has been- 
mentioned that the reunion of the fine capillaries is very slight, 
but here it is quite absent. 

The o-landular epithelium is of the same type as that found in 
Syngnathus, the cells only, as may be seenfrom figure 27, being 
somewhat more columnar in form. The epithelial cells of Gobius 
are relatively large, and, as just sfcated, columnar in form and in 
my preparations faintly granular. They are folded in the simple 
manner already described for Syngnathus and Anguilla, large 
capillaries being situated in the interior of the folds *. In figure 27 
are represented three epithelial folds of the gas glancl, underlying 
which are a few of the multitudinous fine capillaries composing 
the rete mirabile. Even in this figure it can be observed that 
several of the cells contain spherical clear spaces and that two 
small capillaries have encroached on the space previously occupied 
by the cytoplasm of two cells of the gas gland. These intra- 
cellular bubbles and capillaries and the intracellular lumina, which 
1 shall also shortly mention, are cytological fea,tures which have, 
within the last year or so, been studied in great detail by Jaeger 
(44-47), Bykowski &, Nusbaum (24), and Reis & Nusbaum (54, 
55 62-66) and they are of considerable interest from several 
standpoints. I shall discuss their significance later; for the 
present I may remark that I think I am justified in redescribing 
these structures, especially in view of the controversy concerning 
the mode of gas-production considered in Part II. That the fine 
capillaries of the rete mirabile often find their way deep into the 
substance of the glandular cells, as well as between them, is proved 
in Gobius by a veiy moderate amount of inspection under a high 
power of the microscope (PI. IV. figs. 28, 29). It is also noticeable 
that the protoplasm of that border of the glandular cells in contact 
with the capillary develops a dark and often striped appearance, 
which Nusbaum & Eeis suggest is due to difTusion streams of 
nutritive matter passing from the blood into the gland-cells, but 
which I believe is intimately concerned with the production of 
the gas contained in the bladder (see Part II.). Concerning the 
presence of vaciiolar spaces in the cytoplasm of the cells of the 
o-as gland, nobody can doubt their existence who has examined 
well-preserved preparations of gas glands which are in an active 
condition. These vacuolar spaces are with good reason assumed 
to represent bubbles of gas being generated within the cell- 
substance (see Part II. and Appendix B), but it is evident that only 
on occasion will the gas gland be fixed and preserved when active ; 
in the majority of cases, preparations ot gas glands will not 
exhibit these intracellular gas bubbles, simply because the gas 

* Eels (64) states that tlie folds branch and anastomose in G-ohius; in all my 
nrenarat'ions of Gobius nic/er, Gobius paganellus and Gobms minutus (also in 
Gobms ca}rito—Cos:g\, 25) the folds are quite simple, as described in the text. 



GAS GLANDS OF SOME TELEOSTEAX FISIJES. 197 

gland is normally (juit^seent *. To deny the existence of these 
cytoplasmic bubbles simply because cei-tain microscopic pre- 
parations do not show them is illogical, to say the least, since 
intracellular gas babbles, unlike intracellular capillaiies, are not 
pei-manent but transitory structures. Ot" over fifty series of 
preparations which I have made of numerous types of gas gland, 
not more than seven or eight show these intracellular gas bubbles 
in an unmistakable manner. In the type of gas gland under 
consideration, one only of my six or seven series of preparations 
of the gas glands of different specimens of Gohius niger and 
G. paganelliis exhibits a few gas bubbles (fig. 27). I have 
seen these bubbles best in a preparation of Gohius Tninutus (PI. V. 
fig. 35), in which the gas gland cells were fixed in the active con- 
dition. As shown in figure 35, the majority of the gas-producing 
cells possess large vacuolar spaces in their cytoplasm usually present 
in the vicinity of the nucleus ; indeed, the nucleus is often so 
adpressed as to assume a crescentic form. These vacuoles, which, 
as just mentioned, are assumed to repiesent the moulds, so to 
speak, which contained gas bubbles, just as liver- and kidney-cells 
exhibit similar vacuolar spaces which contained liquid globules 
(text-fig. 60, p, 225), are of various sizes, and some can be seen in 
the act of being ejected from the cell substance into the bladder 
lumen, where they are also occasionally found in a liberated 
condition (see Appendix B). The bursting of these gas bubbles 
is doubtless accountable for the masses of granular matter always 
found, when the gland is active, in the bladder lumen and gland 
ducts just external to the glandular epithelium, this granular 
matter, of course, having composed the walls of the bubbles. The 
gland cells, when active, always have that portion of their 
cytoplasm situated next vascular tissue veiy distinct from the 
rest, it being, as already described, very dense and often striated 
in appearance ; the rest of the cj^toplasm usually assumes a 
" stringy " appearance and is very vacuolate f. It may also be 
mentioned that the nucleus is not situated in the dense cytoplasm 
next the blood, in which respect gas gland cells show a mai'ked 
difi"erence from the cells of the pancreas, e. g. 

In addition to intracellular capillaries and gas bubbles there 
are also occasionall}' to be seen intracellular lumina or ducts — 
continuations of the intercellular ducts into the substance of the 
gland cells. These, again, vary greatly in different types of gas 
gland and in different specimens of each type. In none of my 
preparations are they very numerous, and doubtless, like the gas 
bubbles, the smaller intracellular lumina ai-e transitory structures. 
I have figured one or two of these structures in connection with 
the gas gland of Oj^hidmrn harhatnm, a type very effectually 
studied by Reis k, Nusbaum (62), and they are occasionally to be 
found in my preparations of Gohius niger and Gobivs paganellus 
opening into the numerous finer intercellular ducts. Since these 

* See Addenda (D- . . 

t Fov excellent descriptions and figures of the?e gas bubbles in the cytoplasm of 
the sjland cells, see the papers of Hykowski & Nusbanm (24) and Keis & Nusbauni 
(55,62,63). 

Proc. Zool. Soc— 1911. No. XIV. 14 



198 DU. W. N. F. WOODLAND ON THE 

intrn.eelliilai' cliannels liave been so tliorouglily studied hj the 
anthers just named, I shall not refer to them in any detail. I 
will merely say that since, accoi-ding to Jaeger (47), gas glands 
are able, when active, to swell to three times their normal size, 
and that the finer ducts only become visible when filled with 
fiuid, it is not surprising that they are only visible in very few 
preparations. 

One or two other histological features to be observed in serial 
sections of the gas gland and associated structures of Gobius niger 
and other types remain to be mentioned. In transvei'se sections 
across the anterior end of the rete mirabile, and only at its 
anterior end, where the arteries are easily distinguishable from 
the veins, all the ai'teries possess a peculiar endothelium. This 
endothelium (PI. TV. fig. 32) contains many more nuclei than usual, 
and these, instead of being flattened in the usual way, are more or 
less spherical, and, each being contained in a small mass of 
cytoplasm, project into the vascular lumen in the manner shown 
in the figure. This peculiar endothelium, in the case of the 
smaller arterial capillaries, results in such a thickening of the 
wall (and incidentally in some of the smallest capillaries a blocking- 
up of the lumen) as to cause the capillary to bear a strong 
resemblance to a bile- or pancreatic duct ; indeed, were it not for 
the presence of blood corpuscles these small capillaries would be 
almost unrecognisable as such. This type of vascular epithelium, 
which must be well-known to histologists, in all probability results 
from the contraction of the pulsatile arteries in forcing the blood 
through the rete, the narrowed circumference of the endothelium 
causing the cells both to assume a globular form and to protrude 
into the lumen of the vessel. 

Concerning the posterior extension of the difi"use pancreatic 
acini among the arteries and veins anterior to the formation of 
the rete mirabile (fig. 25, e. g.) there is little to note other tha,n 
the fact itself, which I have not seen recorded by writers on the 
teleost pancreas (see list of references to j^ancreas literature 
below). When first studying teleost "red bodies" I was misled 
into supposing that this extraordinary extension of the pancreatic 
acini through and right into the bladder wall in many teleost 
genera rejDresented a new gland specially developed in connection 
with the rete mirabile. I suggested (78) that the purpose of this 
supposed new gland was the abstraction from the venous blood 
stream of the globules and granules resulting from the breaking- 
up of the red blood corpuscles referred to above, and in my sections 
(stained by the picro-indigo-carmine method described in Ap- 
pendix A) the zymogen granules present in the pancreas cells 
often strongly resemble erythrocyte globules which have been ab- 
stracted from the blood stream [cf. text-fig. 56 and PI. III. fig. 21). 
Adopting the view of Jaeger (fully discussed in Part II.) that 
the cells of the gas gland produce a toxin for the purpose of 
breaking up the erythi'ocytes and so enable themselves to absorb 
more easily the oxygen in the blood stream which they sub- 
sequently pump into the gas bladder, I suggested that the purpose 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 



199 



of the rete mirabile — a structure essentially consisting of the 
intimate juxtaposition of the veins returning the blood from the 
gas gland and carrying the hypothetical toxin, with the arteries 
carrj'ing freshly-oxygenated blood to the gas gland — was to allow 
the toxin in its venous capillaries to diffuse into its arterial 
capillaries and so to disintegrate the oxygen-laden erythrocytes in 
time for the oxygen to be available for abstraction by the gas 
glaiid cells by the time the arterial blood reached the gas gland. 
On this view the veins of the rete mirabile would be laden with 
granular matter (as indeed they are — see fig. 32) which requires 
to be eliminated, and I suggested that this pi'ocess of elimination 
was the function of these pancreatic masses which are so closely 
connected with the veins at the anterior end of the rete — the kind 

Text-fig. 56 ( X circ. 470). 




Vein suvroumled by modified acini (tlivee shown) of the pancreas 
in Nerophis eequorius. 



of vessels and the identical position that the hypothesis would 
suggest ; in other words, the hypothesis was supported by the 
position of the gland, the modification of the pancreatic acini 
surrounding the veins (described below), the great similarity 
between the eryttirocyte globules and granules in the blood, the 
zymogen granules in the panci'eatic cells and the granules in the 
pancreatic ducts, and the a i^riori necessity for the elimination of 
the erythrocyte granular matter. This hypothesis, however, con- 
cerning the function of what I subsequently recognized as the 
pancreas is obviously untenable, since the zymogen granules are 
of course present in pancreatic cells not associated with the rete 
mirabile veins and, as in higher Vertebrates, do not occur outside 
the pancreas cells ; also the posterior extension of the teleost 

14* 



200 DR. W N F. WOODLAND OK THE 

pancreas into the bladclei^ wall cannot be supposed to be of any- 
more significance than its penetration into the liver. I failed at 
first to recognize this gland as the pancreas, both because of its 
peculiar position inside the bladder wall and because of the 
peculiar modification of the pancreatic acini in juxtaposition with 
the veins, neither of which peculiarities have been described by 
writers on the teleost pancreas. Another small peculiarity existing 
in several teleost genera (Gobius niger, e. g.) which I studied more 
than others was the syncytial character of the wall of the pancreatic 
duct (PI. V.fig. 33 and Pi". IV. fig. 34), a feature not present in most 
teleosts. The modification of the pancreatic acini in connection ' 
with the veins (veins from the rete mirabile as well as hepa.tic- 
portal veins from the gut wall) just referred to consists of the cells - 
of each acinus next the thin vein wall being drawn out, the cells 
on the side remote from the- vein being quite short, as shown in 
text-fig. 56. This figure also indicates the similarity between the 
zymogen granules and the erythrocyte globules, both globules and 
zymogen granules being stained a bright emerald-green in my 
preparations. As stated in Part II., I now have good reason to 
believe that the erythrocyte granular matter is abstracted from 
the blood in part by the liver and in part by special cell- 
masses situated near the kidneys. Laguesse (4-10), K-ennie 
(17), and others have shown that the teleost pancreas resembles in 
all essentials the pancreas of higher Vertebrates — in the characters 
of the acini, ducts, zymogen granules, and the presence of centro- 
acinal cells and areas of Langerha,ns. With respect to these last, 
I may mention incidentally that I can fully confirm the results of 
Rennie (17) and there can be but little doubt now that these 
areas constitute an organ physiologically and in many teleosts 
anatomically distinct from the pancreatic acini. This view of 
the physiological independence of the islets of Langerhans is sup- 
ported by the researches of Diamare (3), Flint (3 a), Opie (14, 15), 
de Witt (3), Lane (11) and others in opposition to the view that the 
islets are patches of exhausted pancreas, supported by Lewaschevv 
(12), Laguesse (6, 10), and Dale (1); and, as I have already re- 
marked, researches on the teleost pancreas indicate quite plainly 
that the former is the correct view. 

SiPHONOSTOMA TYPHLE (rondeletii Delar.), Nerophis 

^QUORIUS. 

The " red bodies " of these two genera are, like that of S'i/7i- 
gnathus, small red patches situated at the extreme anterior end of 
the bladder and identical in their plan of construction. The only 
particular in which the " red body " of SipJiono stoma differs from 
that of Syngnathus is that in the fonner the cells of the gas gland 
are much larger and the intervening ducts much narrower ; 
indeed, in the posterior portion of the gland the lumina are so 
narrow as not to be seen ver}'' easily. The large capillaries are 
also more circular in transverse section in Siphonostoma than in 
Syngnatlnts. Gas bubbles and intracellular capillaries are in my 
preparations to be found occasionally in the substance of the 



GAS GLAXUS OF SOME TELEOSTEAN FISHES. 201 

epithelial cells, also the cytoplasm in contnct with the capillai-ies 
is darkened in the usual way. A small amount of granular 
material is present in the bladder lumen and gland ducts just 
external to the cells. I mu.st also mention a peculiar class of 
corpuscles found in the blood of my specimen of Siphonosioiiia 
typhle. These peculiar corpu.scles (PI. V. fig.. 36) possess very dense 
cytoplasm, are about half the size of the red corpuscles, and 
contain large nuclei. They are fairly numerous, forming roughly 
about 1 per cent, of the total number of coipuscles. They doubt- 
less i-epresent white corpuscles, though in my preparations they 
to some extent resemble small rounded ganglion cells ; indeed, 
they bear quite a strong resemblance to the large dense ganglion 
cells so often found between the pancrea.tic a,cini, and are in con- 
sequence very conspicuous. I have not observed such conspicuous 
white coi'puscles in the blood of any other of my preparations. 

In Xerojihis the gas gland epithelium is folded like that of 
Gobius niger and possesses the same features. 

Gasterosteus spinachia. 

In this Stickleback the only feature in which the " red body " 
differs from those of the last four genera described is the splitting- 
up of the rete mirabile into three or four bunches supplying the 
gas gland epithelium at different levels ; instead of the initial 
artery and vein wholly dividing up to foim a single I'ete mirabile, 
an artery and a vein are given off from this to form a rete 
mirabile in connection with the most anterior region of the gland 
epithelium, another pair to form a second rete mirabile in con- 
nection with the middle region of the gland epithelium, and, 
finally, the initial artery and vein themselves form a third rete 
mirabile to supply the rest of the gas gland. This condition 
in Gasterosteus is intermediate between that described foi 
S'i/vgnathus and that about to be described for Feristctlms. In 
all other respects the " red body " of Gasterosteus I'esembles the 
Syngnathus type. 

Peristethus cataphracxus {Peristedion catapuractusi 

C. v.). 

The " red body " of Peristethus is star-shaped (PI. Y. fig. 37), 
owing to the fact that the artery and vein which supply the glan- 
dular epithelium, when arriving at the centre of the star-shaped 
" red body,'' break up each into ten vessels. The score of vessels 
thus formed sort themselves out into pairs of vessels, each pair 
consisting of an artery and a vein. These pairs radiate out from 
the point of subdivision of the original artery and vein like the 
spokes of a wheel and give rise to ten elongated retia mirabilia in 
the manner shown in text-fig. 57. Each rete mirabile supplies a 
radially-disposed tract of glandular epithelium developed as usual 
from the innermost cellular layer of the bladder wall. The capil- 
lai-ies of each rete mirabile coalesce to a certain extent to form 
lai-ger vessels before supplying the gas gland (not indicated in 



202 



DR. W. N. F. WOODLANB ON THE 



text-fig. 57), but not to the same extent as in Syngnathus. Figure 
38 (PI. Y.) represents a section (A-B in text-fig. 57) across the 
anterior end of the star-shaped " red body." 

The epithelium of the gas gland is folded as in Gobius niger, 
but, as in Syngnathus, it becomes considerably thickened by the 
elongation and fusion of the folds in the vicinity of the rete 
mirabile. The cells are rather small and short-columnar in form, 
and, in the particular specimen which I examined, contained 
numerous gas bubbles (figs. 39, 40, 41) ; in other Avords, the gland 
-was here in a highly active condition when preser\ed. 






um 

The construction of tlie " red body " of Feristetlnts cataphrcictus. 

E., epithelium of gas gland; R.M., rete mirabile. Fig. 38 (PL V.) represents a 
section taken across A-B. 

The most remarkable feature about the " red body " of 7^eri- 
steihus is, of course, the radial disposition of the divisions of the rete 
mirabile and the glandular epithelium ; in every other respect it 
appertains to the SyngnatJms type as above defined. 

Trigla hirundo (corax Bp.). 

The bladder of Trigla hirundo, when observed from the ventral 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 203 

aspect, appears as an elongated sac, the vcnti-al wall of which is 
very muscular in the median line posteriorly. At the sides of this 
muscuhir posterior median portion of the bladder wmII there are 
to be seen two ]'ed streaks, which are the lateiul divisions of the 
" red body" (PI. VI. fig. 42). In a series of transverse sections it 
can be seen that the cavity of the bladder has a peculiar conform- 
ation* ; anteriorly the bladder cavity consists of two laterally-pliiced 
channels, which end blindly anteriorly and proceed posteriorly 
over a distance equal to about one-third the length of the bladder 
when each divides into two ; these four chaniiels thus formed then 
extend to about midway in the length of the bladder, when the 
two inner channels coalesce to form a wide median cluuuber, 
which, with the tAvo external channels, extends to the posterior ex- 
tremity of the bladder (PI. Y. fig. 43). It will be observed in figure 
43 that two great median bands of muscle (M.R.) are developed 
in the ventral wall of the posterior half of the bladder, and this 
explains why it is that only the lateral "red bodies" are seen 
when the bladder is viewed from the ventral aspect (fig. 43). 
Also shown in this figure are the two laterally-placed sti'ands (S) 
of nerve-fibres developed in connection with the two muscle- 
bands. 

The epithelium of the gas gland is folded in the manner cha- 
ractei-istic of this type of " red body," but in Triyla it is more 
folded than in previous examples (PI. VI. fig. 44), the primary 
folds bearing small lateral folds. In some places the depth of the 
folding is increased as in Syngnathus. The cells are of moderate 
size and columnar in form. The rete mirabile is divided u[) into 
capillary tufts, i. e. there are numerous small retia mirabilia which 
supply the epithelium at intervals in its course ; in other words, 
pairs of vessels are given off at intervals from the bladdei' ai-tery 
and vein, each pair consisting of an artery and a vein and forming 
a rete mirabile to supply a certain area of gas gland. 

TrigLiA gurnardus. 

Concerning the gas gland epithelium of this species, see p. 207, 
below, 

Smaris maurii, Smaris vulgaris. 

The " red body " of Smaris vidgaris has the macroscopic appear- 
ance shown in figure 45 (PI. VI.). Figure 46 (PL V.) represents 
diagi'ammatically a transverse section aci'oss the anterior end of 
the " red body," the primary artery and vein entering at the 
centre of the oval mass. 

Concerning the character of the gas gland epithelium of these 
two species (PI. VI. fig. 47), see p. 207, below. 

Previoihsly described " Red Bodies " of the Syngnathus l^yj^e. 

The glandular epithelia of the "red bodies" of Gohins capilo, 
Trigla gurnardiis (see my description below), and Gasterosteus 

* Sue descriptiun in Mlliiu-Edwardb (49). 



204 DR. W. N. F. WOODLAND ON THE 

actdeatus have been described and figured by Coggi (25) ; Gaster- 
osieus aculeatus has also been described and figured by Deineka 
(29) ; Blennms, Gobius, Syngnathus, and other types have been 
briefly described by Reis (64). All these " red bodies" evidently 
belong to the Syngnathus type as defined above. 

A Third Type of Bladder Vasculo-glandular Apparatus, 
as exemplified in Ophidium barbatum L. 

The " red body " of Ophidium barbatum has received a con- 
siderable amount of attention during the last few years, especially 
from the cytological standpoint (Bykowski & Nusbaum, 24 and 
Reis & Nusbaum, 62, 63), since the huge cells composing the gas 
gland offer exceptional facilities for enquiry in this direction. 
But, as in some other instances, good figures of the gross anatomy 
do not exist, so far as I am aware, and hence I shall make no 
apology for recapitulating already-known facts in a manner that 
shall be intelligible to the reader who has not worked at the 
subject. 

The " red body '' of Ophidium barbatum is situated anteriorly 
in the ventral wall of the bladder and has the curious appearance 
shown in figure 48(P1. YL). The fan-shaped deep red portion is, of 
course, the rete mirabile, which is here an undivided body, and it is 
formed by an artery and vein which enter at the pointed posterior 
apex. The pink margin surrounding the anterior border of the rete 
is, obviously, the gas gland, A transverse section i-unning mid- 
way through the " red body " exhibits the parts diagrammatically 
indicated in figure 49 (the outer layers of the bladder have been 
stripped off). In the centre is the rete mirabile and at the sides 
are the enormously thiciiened glandular portions of the (elsewhere) 
flattened lining epithelium. As in some former instances, there is 
no reunion of the capillaries of the rete mirabile before supplying 
the glandular epithelium : they directly supply the gas gland. 

Kow in this type of " red body "' the gas gland does not consist 
of a single layer of cells which has become folded, but of a many- 
layered mass of cells, the arrangement of which bears no resem- 
blance to the folded condition of the single-layered epithelium of 
the " red-bodies " already described (see figs. 54, 58, 63 for 
examples of the massive tj^pe of glnndj, and, judging from the 
development of this massive type of gas gland epithelium as seen 
in Atherina, it never does, at any stage of its development, bear a 
resemblance to the folded type of gland. The many-layered or 
massive kind of gas gland epithelium, as seen in Ophidiimi, 
Atheri7ia, and other genera to be mentioned, is, then, a distinct 
type ; the question raised bv Reis in a recent paper (64) as to 
whether there exist forms of bladder epithelium transitional be- 
tween these two types is one which I shall discuss when describing 
Atherina. The faintly-granular cells composing this m.any-layered 
gas gland of OphidAum are of quite moderate dimensions in the 
vicinity of the rete mirabile, but become larger the further they 
are removed, and the cells at the remote edge of the gland are 
remarkable for their large size, In this type of gland there exist 



\ 



GAS GLANDS OK SOME TIXHOSTKAN FISHES. 205 

no l;n-ge duets compai'a1)]e with those we have seen in the glands 
consisting of folded epitlielium ; on the contrary, the only ducts 
visible are narrow slit-like channels occasionally to be detected in 
their coarse between the cells. These narrow channels vaiy in 
width greatly according to the state of activity of the gland ; they 
are necessarily numerous (though by no means always to be 
observed) in order to provide means of exit to the surface for all 
the numerous cells concerned, and they open on the gland surface 
inside the bladder lumen by small pores. Also penetrating the 
mass of the gland are numerous fine capillaries supplying the 
individual cells with nuti'iment a,nd oxygen on the sides remote 
from those facing the ducts just mentioned. All the intracellular 
structures previously mentioned as occurring in connection with 
cells of the gas gland are here shown in a very obvious manner. 
Figure 50 (PI. VI.) shows intrncellularlumina. one in longitudinal 
section. As to the significance of these intracellular passages found 
in gas gland cells, it is not necessary to regard them all as possessing 
purely a nutritive utility as ISTusbaum & E.eis have supposed ; it 
seems more probable, judging from their appearance in such cases 
as that illustrated in figure 50, that some at least are concerned 
with the passage of the gas bubble from the interioi- of the cell 
to the exterior; they are veiy rare in their occurrence in my 
preparations and in transverse section closely resemble bubbles, 
especially if near the nucleus. The whole mass of cells composing 
the gas gland is therefore penetrated by capillaries and gland 
liimina. I have not obsei-ved a pancieas in the region of the 
rete in Ojyhidiitm. 

Other Examples of the Third or Ophidium Type of " Red Bodyr 

Box Boops L. 

The "red body" of Box looj^s has the curious macroscopic 
appearance shown in figure 51 (PI. VI.). Figure 52 (PI. VIL)isa 
diagram of a transverse section through the middle of the " i"ed 
body," and the appearance of two concentric loops is thus seen to be 
due to the disposition of the rete mirabile. The primary artery 
and vein enter the " red body " postei'iorly and break up into 
smaller ai"teries and veins (interiningkd with pancreatic acini) to 
form the rete mirabile as usual. The glandular epithelium consists, 
as in Ojyhidium, of a mass of cells interpenetrated by capillaries 
and lumina, but here the cells are all approximately equal in size 
and the gas gland is more than a dozen cells thick. 

Atherina hepsetus L. 

In macroscopic apjoearance the " red hoAj" oi Atherina hep)seius 
is diffuse in form (PI. VI. fig. 53) and consists of tufts of retia 
mirabilia borne laterally on a median pair of vessels (artery and 
vein) with cori'esponding tracts of glandular epithelium . The glan- 
dular epithelium (PI. VII. fig. 54) is very similar to that of Box 
hoops ; in other words, it consists in its thickest parts of amass of 
small cells, seven or eight cells in thickness, with inter- and 



206 DR. W. N. F. WOODLAND ON TUE 

occasional intra-cellular ducts and ca])illaries. I have not observtd 
a pancreas in the region of tlie I'ete. The cell-outlines of the 
gas gland are faint and a small amount of giunular material is 
present in the bladder lumen in my preparation. 

1 stated above that the massive type of gas gland is quite 
distinct from the folded type in certain genera, and, indeed, these 
two types of bladder ej^ithelium are almost always easily distin- 
guishable fi'om each other in the genera which I have studied : 
genera, with a few exceptions to be mentioned, either possess 
the folded or the massive type of gland and not glands inter- 
mediate in structure. In young specimens of Atlterina about 
15 mm. in length, that portion of the bbulder epithelium which 
is about to produce the gas gland is at fiist composed of a single 
layer of cubica,l or short cokunnar cells, and it is worthy of notice 
that these cells, in order to build up the fully-formed massive 
gland, all divide at right angles to the plane of the bladder wall 
(PI, VII. fig. 55), i. e. the successive planes of cell-cleavage are 
parallel with the plane of the bladder wall — the initial unilaminar 
epithelium never becoming folded at any stage of development. 
As the gas gland cells multiply connective tissue cells may be 
observed to extend in between them in order to furnish the 
connective and vascular tissues which penetrate tlie mass of the 
gland when fully formed. Also if figures 54 and 55 be compared, 
it will be seen that the cells (especially the nuclei) of the gas 
gland of the adult Atherina are noticeably smaller as compaied 
with those in the young tish. The fact that there is no indication 
of folding of the bladder epithelium during development of the 
massive type of gland is important, since Reis (64) supposes, on 
the ground that there exist forms of bladder epithelium more 
or less intermediate in structure between the folded and massive 
types, that the latter has been derived from the former, and if 
such be the case we might expect that the massive type of gland in 
its development would pass through a folded developmental stage. 
As we have seen, this is not the case in Atherina and presumably 
not in other genera possessing typically massive glands. I assume 
this with the more confidence since in some gas glands which, 
owing to the exceedingly close apposition of the folds, at fiist 
sight appear to be massive, it is always possible to refer them 
at oncG to the folded type by observing in each such case the 
margins of the gland where it merges into the squamous epithe- 
lium lining the greater part of the bladder cavity, since here the 
folds are quite simple : such marginal folds arc never present in 
true massive glands. 

Reis, in contending that the massive type of gland is but a 
derivative of the folded type, instances as transitional forms 
between the two types the gas gland epithelia of Syvgnathus and 
Girardinus. I have not examined GirardimLS, but the gas gland 
epithelium of my specimens of Syngnathus can certainly not be 
said to be transitional, since, as my description above of this epithe- 
lium clearly shows, the massiveness of the gland merely results from 
the extreme leneth of the folds ^nd their occasional anastom^osis : 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 207 

the glandular epithelium is always unilaniinav and the folds always 
clearly recognizable. My prepn.rations do not at all confirm 
Rei.s's statement that the tabular outgrowths are so numerous nt 
the base of the gland that they lose their lumina through mutual 
contact, and form almost complete layers of epithelial cells. On 
the other hand, the gas glands of some other genera which 1 have 
examined certainly do appear to be transitional in structure 
between the folded and massive types, viz. those of Trigla gurnar- 
dus, Smaris maurii, and ASmaris indgaris. I may say at once 
that in all three species, especially Triyla guriutrdus and Smaris 
maiM'ii, the gas gland shoAvs distinct signs of being of the folded 
type, and in the two species just mentioned the folds are quite 
distinct and unmodified at the edges of the glands, but, on the 
other hand, the surface of the gland is covered by a continuous 
single layer of columnar epithelium (never present in the typical 
folded gland) and throughout the mass of the gland the cells at 
certain points lose their unilaminar arrangement *. In Trigla 
gihrnardus f", and perhaps also in the other two species, the explana- 
tion of the superficial cell-layer seems to be that at the edges of 
the gland the unilaminar epithelium splits into two layers, the 
upper remaining unfolded and forming the superficial cell-layer 
and the lower becoming folded in the usual manner (text-fig. 58), 
the folds, however, anastomosing to a considerable extent and, as 

Text-fig. 58. 




Diagram to explain the probable construction of the gas glands of 
Trigla gurnardiis and Smaris inaurii. 

already mentioned, the cells in places becoming clustered into 
groups more than one layer in thickness. In Smaris maurii (PI. VI. 
fig. 47) the individual cells are larger than in Trigla gurnardus 
and the anastomosing folds more closely packed ; here and there 
the cells are several layers thick, but in general they are arranged 
in single folded rows as in Trigla gurnardtts. In Smaris vulgaris 
the epithelium has almost entirely lost its folded character, though 
this is sometimes to be detected at the edges of the gland. Thus 
the gas glands of these three species t are to a certain extent 

* In many massive glands the cells are often i)i places arranged in single rows, 
and this is evidently due to the necessity (which Opi)el points out : see footnote on 
page 208) of each cell being in contact on one side with a duct and on the other 
with a blood vessel. 

t Coggi (25) provides a figure of the epithelium of this species showing a simple 
folded character; the epithelium in my specimens more resembles his figure of the 
epithelium of Ilotella mediterranea with the addition of a superficial covering- 
layer of columnar cells. 

X And possibly those of Folyacantlnis- (Mavrnpodus) des-cribed by lleis & 
Xusbaum (62) and MoteUa described by Coggi (25). 



208 DR. W. N. F. WOODLAND ON THE 

intermediate in character between the folded and massive types, 
but we may still retain these two categories, since we have reason 
to sujjpose that all of these three supposed transitional forms of gas 
gland really belong to the folded type and that they have only 
secondarily acquired features characteristic of the massive type 
by the splitting-ofF of a superficial cell-layer covering the surface 
of the gland, by close apposition and anastomosis of the elongated 
folds, and by occasional local prolifei-ation of the cells ; in other 
words, until their development is known, we may provisionally 
distinguish between truly or primarily massive glands which 
never exhibit the folded arrangement of cells at any stage of 
development, and ps-eudo- or secondarily-massive glands which are 
secondarily derived from the folded type*. To me Reis's sugges- 
tion that all massive glands have been derived from folded glands 
seems very improbable, since on this view the folded condition 
should occur as a stage in their development, and we have seen 
that this is not the case in at least one typical genus {Atherina). 
It seems more probable that both folded and massive types have 
originated independently from the simple unilaminar condition 
which must in all cases form the starting-point of development 
both in ontogeny and phylogeny. 

CoRis JULis t (vulgaris Flem.), 

The " red body " of Coris julis has the circular shape seen in 
figure 56 (PI. VI.), and the rete mirabile supplying the gland is 
noteworthy for the way in which it is broken up into small divisions 
(PL VII. fig. 57). The glandular epithelium (fig. 58), which is 

* Reis's classification of gas gland epithelia (64) seems to me confinnatovy of 
this view. Her classification is as follows : (1) entireij' nnilaminar epithelium 
folded into simple tubular outpushings ; (2) the epithelial folds are umcli folded 
and branched [Keis has made the mistake of including in this group the gas gland 
epithelium of Coi-vina ; in mj' preparations of Corvina niyra the epithelium is of the 
most_ typical massive type] ; (3) the epithelium is in parts folded and in parts 
multilaminar [I cannot agree to the inclusion of Sijngnathus in this category. I 
have not examined Reis's other example — Hipponampits] ; (4) the compact or massive 
glands of Sargtis, Charax, and other genera. Reis regards these four types as a 
developmental series — a mistake I have corrected in the text. Reis also remarks 
that only in the first three types are the ducts obviously the spaces between the 
epithelial folds or tubular outgrowths ; in the fourth there exist no such intertubular 
ducts, intercellular crack-like channels only being present. This last statement 
seems to me to be confirmatory of my contention. 

Reis makes another statement which, as Oppel (58) points out, contravenes all 
probability. This statement is that whereas in the folded type of gland the 
individual gland cell only presents one surface towards the duct lumen (the others 
being contiguous with other cells and with a blood-vessel), in the massive gland the 
individual gland cell presents all its surfaces towards the intercellular lumina pene- 
trating the mass of the gland, i. e. the intercellular lumina, according to Reis, 
surround each cell. Reis suggests that this supposed feature is for the purpose of 
facilitating the exit of the gas through the gland into the bladder. As Oppel 
justly I'emarks, how can a cell thus shut off from all vascular tissue obtain nourish- 
ment ? So far as my observations go, there is no reason to suppose that a single cell 
of the massive gland is not in contact with a capillary. 1 cannot, however, agree 
with Oppel's groundless suggestion that tubular and massive glands possess difierent 
functions, tubular glands being secretory and massive absorptive. Jf this be so, from 
whence do fish with massive glands obtain their bladder oxygen ? 

t Mr. Tate Regan informs me that, according to Holt, " Coris giofrcdi " is merely 
the female form of C.Jtilis. 



GAS GLAXDS OF SOME TELEOSTEAN FISHES. 209 

about twelve cells deep, is nu example of the massive tyjie 
possessing a stiuamous lining epithelium. The lumina consist of 
small crack-like spaces visible here and there between the cells 
(largely arranged in vertical rows) in ti-ansverse sections 
of the gland, and they form a system of fine anastomosing 
channels opening at A^arious points on the surface into the 
bladder lumen. Gas bubbles were occasionally to be found in 
the gland cells. I did not observe any intracellular capil- 
laries, but they doubtless occur; intercellular capillaries, on the 
other hand, are extremely plentiful and occur largely near the 
extreme edge of the glaud underlying the layer of flattened 
cells. The glandular epithelium of Coris julis difiers from 
preceding examples of the massive type in that the cells lying 
next the bladder cavity foim the squamous layer, or in places 
layers, just mentioned. 

It was doubtless because Corning (28) chiefly examined " red 
bodies " of this extreme massive type, as e. y. that of the Pike, 
that he concluded that the bladder glands possess no lumina, and, 
indeed, were it not for our knowledge that most bladder glands 
possess lumina and that these vary greatly in volume according to 
the state of activity of the gland, we should pi'obably come to the 
conclusion if studying exclusively the same mateiial. 

CORVINA NIGRA Cuv., SaeGUS RONBELETIl C. V., BaLISTES 

CAPRiscus L., Caranx trachurus and Serranus cabrilla L. 

The "red bodies "of these genera all belong to the massive 
epithelium type, although they vary in a^ppearance macroscopically 
(Pis. VII. (fc VIII. ligs. 59-61) in correspondence with the vaiying 
configuration of the rete mirabile. The " red body " of Batistes 
alone is situated near the posterior end of the ventiul wall of the 
bladder. In all cases the glandular epithelium has the usual 
structure — a thick mass of cells, penetrated by large or small 
intercellular and intracellular lumina and capillaries. 

Zeus faber L. 

The " red body " of Zeus faber is of the curious form depicted in 
figure 62 (PI. VIII.), and occupies the anterior half of the ventral 
surface of the bladder wall. It has already been figured by Vincent 
& Barnes (75). The " red body " consists of two lateral divisions, 
each division being shaped like a C divided into three parts, and 
its concavity facing that of its companion. The primary artery 
and vein enter at about the centre of the " red body " and supply 
branches to the retia mirabilia lying on the inner sides of the gas 
gland thickenings. Anteriorly to the " i-ed body " a thick median 
longitudinal muscle- band is developed in the ventral bladder wall. 
The glandular epithelium of the John Dory is of the ordinary 
massive type, though more folded in appearance than is usual. 
It is significant that Vincent & Barnes, who wished to reduce 
all " red body " glandular epithelia to the folded type, remark 



210 DR. W. N. F. WOODLAND ON THE 

that " we have not been able to obtain sections which show the 
direct transition to the tubular glands, but there is no doubt that 
the epithelium takes on fairly suddenly a markedly glandular 
aspect." 

A conspicuous feature in the gas gland of Zeus is the presence, 
among cells of the ordinary size, of giant cells and nuclei six to 
ten tiroes as large (PI. IX. tig. 65). Similar giant cells also occur in 
Ophidium harhatimn, Fierasfer acus, Peristethus cataphr actus ^ 
Balistes capriscus, Trigla gnrnardus, and, in some instances, in 
Gohius paganellus and in other genera. The only suggestion 
concerning the origin of these giant cells with which I am 
acquainted is that implied by Deineka (29), who states that the 
similar giant cells in Perca give rise by amitotic division of the 
niocleits to groups of cells of the usual size — the implication being 
that the giant cells are primary products of development and cells 
of normal size secondary products. My observations have led me 
to a somewhat different conclusion. In the first place, in many 
gas glands containing giant cells {Zeus, e. g.), these giant cells 
usually lie more or less remote from the region where the blood- 
vessels enter the gland — giant cells are nearly always found 
towards the periphery or edge of the gland epithelium ; secondly, 
there is every gradation in size from the smallest to the giant 
gas gland cells, the smallest always being situated in those 
portions of the gland next the large blood vessels, i. e. at the bases 
of the folds of folded glands and at the " hub " or point of entry of 
the blood-vessels in massive glands (see figs. 46, 49, 52, on Plates Y., 
VI., YII. e.g.) in addition to other regions, the cells of intermediate 
size like the smallest cells being found in most regions of the gland 
and the giant cells, as already mentioned, towards the peripheral 
portions of the gland * ; thirdly, among the smallest cells mitotic 
figures are quite common in many of my prepai-ations, but among 
the intermediate and giant cells it is rare to find mitosis, though 
I have seen several cases of it in intermediate cells and once or 
twice even in cells approaching giant size ; finally, I can confirm 
Deineka in his statement that the great majority of giant cells and 
many intermediate cells divide amitotically. Figure 63 (PI. VIII.) 
shows the appearance of the ej^ithelium at the base of the gland of 
Zeus, where the capillaries of the rete enter. It will be observed 
that in this genus many of the basal portions of the massive 
epithelium are syncytial in character, no cell-outlines being distin- 
guishable, and that occasionally mitotic figures are present among 
the nviclei of these syncytia. As we proceed from these syncytial 
masses towards the periphery of the gland, cell-outlines soon become 
more and more distinct and larger in size, but, as we might 
anticipate, though the giant cells are usually found towards the 
periphery of the gland, j^et giant nuclei and nuclei of a size 
intermediate between these and the smallest nuclei are to be found 

* Reis remarks that in Sao'ffus (in wliicli there are no giant cells) the cells are' 
largest next the blood-vessels and decrease in size towards the periphery', where they 
are squamous in chai'acter. This is not the case in iny preparations of Sarcjus 
rondeletii, where the cells are practically uniform in size at all depths of the gland, 
except perhaps at the ^'ery edge, where they are squamous. 



GAS GLANDS OF SOAIE TELEOSTEAX FISUES. 211 

in the Kynoytial masses at tlio base of tlie gland. In cells of 
intermediate size and in giant cells ninitosis occurs, as I have 
already state 1. Figure 64 illustrates several phases of direct 
division, l)ut L must also mention that the huge nuclei of many 
giant cells, like those of myeloplaxes of bone-marrow, appear 
sometimes to divide into several — three, four, or more — nuclear 
lumps at a single division, though I do not remember to have 
seen cells containing more than two nuclei in the resting condition. 
Tiiese various fa.cts considered collectively seem to suggest that in 
cases like Zeus the initial unilaminar epithelium of the bladder 
wall must contain nuclei of various sizes and that the nuclei of 
each size, associated with cytoplasm, multiply in order to produce 
the massive gland of the adult fish. As the nuclei become more 
numerous they migrate from the centime of proliferation, the small 
nuclei occupying cytoplasm vvhich, owing to the size of the nuclei, 
becomes subdivided up into small areas, the lai-ger nuclei, fewer 
in number, also occupying cytoplasm which, owing to the superior 
size of the nuclei and their slow rate of division, remains to a 
large extent unsubdivided, so forming the giant cells round the 
giant nuclei. There is no evidence in my preparations of giant 
cells splitting up into groups of small cells. The small cells 
multiply by mitotic division, and amitosis is only found in the 
case of the larger nuclei — those of the giant cells and larger 
intermediate cells. As to the origin of the giant nuclei I can 
give no infoi-mation. It is stated that the cells of bone- marrow 
and some glands divide mitotically after undergoing amitosis, so 
that there is no theoretical objection to supposing that the small 
syncytial niiclei have been produced, as Deineka supposes, by 
amitotic subdivision of giant nuclei, but I have never seen any 
appearance of this taking place and I very much doubt its 
occuri-ence : nearly all the appeai'ances of amitosis which I have 
observed have been in connection with the nuclei of giant cells, 
i. e. far away from the syncytia. As to the reason for amitosis 
occurring in connection with giant nuclei, it is only possible to 
readvance the old suggestion that it is a prelude to degeneration. 
The gas gland is in Zeus and presumably other types constantly 
being regenerated at its base, and therefore there is ground for 
supposing that cells at the periphery of the gland become used up 
and degeneration is the natural result of this. But this suj-)pose<l 
degeneration of the cells has nothing to do with the violent 
decomposition of cell-substance assumed to occui' by Nusliaum &. 
Reis in connection with the production of gas — a phenomenon for 
the existence of which I can find no evidence and in which I 
fpiite disbelieve. 

Gadus morrhua. 

The "red body"' of the Cod* is knoAvn to every student of 

comparative anatom}' as a large red mass situated antei'iorly in the 

bladder. As Vincent k Barnes (75) say, the free surface of the 

" red gland '' shows " many small clubbed processes, packed 

* I used common "CoiUing'' caught off Deal. 



212 DR. W. N. F. WOODLAND OX THE 

tightly together, each consisting of a red mass of tissue, sur- 
mounted by a cap of yellowish mucilaginous-looking material." 
Each club-shaped process " consists essentially of two parts : — 
(1) a mass of capillary blood-vessels, lying side by side, conveying 
blood to and taking blood from the second parts ; (2) a glandular 
portion." In figure 66 (Pi. YIII.) is represented diagrammatically 
a transverse section of the gland of the "Codling," which shows 
the " clubbed processes " well in section, also one " cap of 
mucilaginous-looking " connective tissue. Vincent & Barnes go on 
to say that " when the capillaries arrive at a point about "2 mnii 
from the free surface of the gland, they interdigitate with 
involutions of the glandular epithelium, which dips down between 
them and forms a covering for them, thus constituting tubular 
glands "2 mm. in length." In other words, Vincent &, Barnes 
state that the glandular epithelium of Gaelics is that which I have 
described above for the Eel and SyngiiatJms, and they provide a 
diap-ram of what they suppose to be tlie folded arrangement of the 
epithelium. I am quite willing to admit that the small columnar 
cells which compose the glandular epithelium are in geneiul 
arrano-ed round strands of vascular connective tissue, and that 
occasionally the intercellular lumen is so extensive and so disposed 
as to give the glandular tissue an almost folded appearance, but 
there is no real ground for comparing the massive structure of the 
glandular epithelium of the "Codling" to the folded epithelium 
of Syngnathus, The gas gland of the " Codling " is of the massive 
type, in which the intercellular lumina are very abundant and 
occasionally very large. The capillaries are very small and run 
in the numerous thick strands of connective tissue. As Vincent 
& Barnes mention, caps of connective tissue, continuous with 
the connective tissue penetrating the glandular epithelium, are 
often present. I have not examined them in detail. 

Cepola rubescens L. 

The " red body " of Cepola t^iobescens is situated anteriorly in 
the ventral wall of the bladder and has somewhat the shape of a bell 
(PL VIII. fig. 67), the handle being formed by the rete mirabile. 
The arteiy and vein enter the " red body " anteriorly at the top of 
the bell-handle, and there give rise to the rete mirabile in the usual 
way. The rete mirabile gradually diminishes as it supplies the 
mass of the gas gland, which first appears, in the serifS of trans- 
verse sections, on the right side of the " red body" (text-fig. 59, 
G.E.). The peculiar feature about the " red body " of Gej^ola 
is the curious folding of the glandular mass anteriorly. The gas 
gland extends a considerable way posteriorly, as seen in figure 67, 
and the best way to comprehend the folding anteriorly — the 
foldino-, that is to say, of the whole mass of the gas gland, not 
the folding of the epithelium hitherto described — is to observe the 
diagrams of the sections contained in text-figure 59 in the reverse 
order i. e. observe H first and A last. Diagram H represents the 
olandular epithelium (the dotted area G.E.) as a mass of cells 
developed from the general squamous epithelium {F.E.) lining the 



GAS GLANDS OF SOME TELEOSTEAX FISHES. 



213 



bladder cavity (B.C.) in the median line of the ventral wall of the 
bladder. In diagram G it is to be observed that the squamous 
epithelium on each side of the glandular mass has become folded 
(F.F.E.) and projects into the bladder cavity towards the median 
line. In diagram F these folds have nearly met in the middle 
line and lie dorsal to the gas gland in the bladder cavity ; also it 
will be observed that on the left side the glandular mass itself has 
become completely reflected towards the median line [F.L.G.G.), 
anil lies in close apposition with the rest of the glandular mass. 
Still more anteriorly, the two lateral folds of squamous epithelium 
have, as shown in diagram E, united in the median line, so that 

Text-flir. 59. 




Diagrammatic transverse sections through the anterior end of fig. 67 (PI. VIII.), 
where the rete mirabile supplies the epithelium of tlie gas gland. The diagram 
is full}' explained in the text. 

two separate epithelia now lie in the bladder cavity above the gas 
gland, the upper of which (I.F.E.) is continuous with the squamous 
epithelium lining the rest of the bladder wall, and the lower 
{O.F.E.) is continuous with the two sides (the left reflected as 
just described) of the glandular mass, which latter therefore now 
forms the thickened ventral wall of a tube. Diagram D shows 
that the dorsal thin wall of the tube just mentioned is being 
replaced by extensions of the glandvxlar mass, this replacement of 
the thin epithelium being chiefly due to the increase in size of the 
Proc. Zool. Soc— 1911, No. XY. 15 



214 DR. W. X. F. WOODLAXD OX THE 

reflected left side of the glandular mass. In diagi'am C the 
reflected left side of the gas gland has inci-eased so as to extend 
over the opposite or right side of the gland, a narrow crack 
(continuous posteriorly with the bladder cavity) only being left 
l)etween the dorsal and ventral halves of the gas gland. In 
diagrams B and A, owing to the coalescence of the glandular 
masses of the dorsal and ventral portions of the gas gland on the 
right side {i. e. the replacement of the short strip of thin epithelium 
which in C closed the tube on the right side), the narrow crack 
(C) becomes bounded on all sides l)y the glandular substance, and 
this crack narrows anteriorly until it disappears shortly before 
the gas gland itself terminates. 

The mass of glandular epithelium (PL IX. fig. 68) resembles that 
of Coris julis in being (apparently) almost devoid of gland lumina. 
Small intercellular cracks are visible here and there between the 
cells in my preparations, but that is all to be observed in the inactive 
gland. On the other hand, the glandular mass is penetrated by 
capillaries, and the large cells appear to arrange themselves round 
these capillaries to some extent, giving the gland in section almost 
a lobulate appearance. The cytoplasm of the cells is conspicuously 
darkened where in contact with the capillaries. Now and again 
large cells are found with a very large nucleus, but these are rare. 
Finally, it may be mentioned that the capillaries of the rete 
rairabile possess the peculiar endothelium which I have already 
described in Gobius pagcmellus and found in most " red bodies." 

Pehca fluviatilis. 

The " red body " of the freshwater Perch is a diiTuse organ 
situated anteriorly on the ventral and lateral portions of the 
bladder wall (PI. IX. fig. 69). The retia mirabilia ai-e seen as small 
fan-shaped tufts present at the extremities of the variousbranchings 
of the closely-associated artery and vein (both included in the 
red streak seen in the bladder wall), and bordering these tufts of 
rete are the areas of gas gland. The gas gland is of the ordinary 
massive type, but feebly developed compared with the gas glands 
of most marine fish, consisting in its thickest parts of not more 
than six cell-layers (see text-fig. 62, p. 240) and dwindling to one 
a.t its edges (PI. IX. fig. 70). The giant cells described by Deineka 
in young Perch are only of very moderate size in my preparations *, 
indeed, not so large as those I have figured in the case of Zeus ; as 
in Zeits, amitosis occurs in connection with most of the cells. 
With reference to the suggestion of Deineka that these giant cells 
subdivide into groups of the smaller cells as a consequence of the 
fragmentation of the nucleus, I have already advanced reasons for 
regarding this as improbable (see Zetts). 

Previous Desanptions of the Structure of the Ophidium 
or Third Type of Red Body r 

I shall here merely enumerate the chief histological descriptions 
of gas glands of the massive type. Emery 1880 (33), Fierasfer ; 
* My Perch were from 10-14 cm. in length. 



GAS GLANDS OF SOME TELEOSTEAN FISUES. 215 

Coggi 1886 (25), Serranits, Perca, Ballsfes, Corvina, Julis, Motella 
and other types, ami 1889 (23) Esox lucius ; Corning 1888 (28), 
Esox, Perca and other types; Vincent &, Barnes 1896 (75), 
Zeus, Gaelics and other types; de 8eabra 1897 (31), Esox and others ; 
Jaeger 1903 (44), Sci(ena, Lacioperea; Deineka 1904 (29), Perca, 
Esox and other types; Bykowski & Nusbaum 1904 (24), 
Fierasfer, Ophidium; Reis '& Nusbaum 1905, 1906 (62, 63), 
Poll/acanthus {Macropodus), Flerasfeyr, Ophidium, Gharax, Perca-, 
Reis 1906(64, 65), Sarcjus, Gharax, Pagellus, Gorvina, Umhrina, 
Chri/sophr>/s and others described in a general way. 

Summary of Part I. and General Gonclusions resjjecting 
^ Gas Gland Structure. 

From the foregoing it is now manifest that all teleostean " led 
boilies," *". e. the vasculo-glandular gas-producing organs situated 
in the bladder wall, are composite structures (consisting of two 
distinct and sometimes separate parts — the glandular epithelium 
or gas gland* and the rete mirabilef. The mutual relationships 
of these two parts are quite constant, and though the gas gland 
must be described as the essential part, yet the rete, on account 
of its constant J presence and undoubtedly highly important 
function, is probably of almost equal importance. The rete 
mirabile bipolare geminum (Miiller, 53), i. e. the arterial and 
venous rete already described, is always formed in connection 
with the artery and vein supplying the gas gland, and may either 
be anatomically quite distinct from the glandular epithelium (Eel 
type), the arterial and venous capillaries reuniting to a varying 
extent on the gland side of the rete to form large arteries and 
veins before supplying the glandvilar epithelium, or the rete 
mirabile may be contiguous with the glandular epithelium, the 
capillaries of the rete undergoing little (e. g. Si/7ignatkus) or no 
(e, g. Ophidium) reunion before supplying this. 

The diffuse pancreas, when present, is often intermingled with 
the arteries and veins situated at the extreme anterior end of the 
rete, and so actuall}^ becomes situated inside the bladder wall ; it 
was at first, on account of several j)eculiarities, mistaken by me 
for a new gland with a special function coimected with the gas 
gland, but this view was quite a mistaken one, the penetration 
of the pancreas into the bladder wall being of no physiological 
significance. 

The gross anatomical difterences, often visible to the naked 
eye, which are to be found in " red bodies " taken from different 
fishes, are determined by the disposition and degree of subdivision 
of the rete mirabile — since the distribution of the glandular 

* The " driisige siiuine " of Miiller .and the " corpo epitheli.ile " of Coggi. 

t The " vaso-gaiiglioH " of Owen and " organo vascolare " of Emery. 

X That is to sa}', using the term " gas gland " in the usual sense as equivalent to 
"oxygen gland" (see Part II.), the rete is constantly associated with it; the rete, 
however, is absent in the cases of those bladders the lining-epithelia of which only, 
or principallv, " secrete " nitrogen and carbon dioxide. 

15* 



216 DR. W. N. F. WOODLAND ON THE 

epithelium must evidently correspond with that of the rete 
mirabile. It need hardly be pointed out that the gross ana- 
tomical simplicity or complexity of the "red body" by no means 
always corresponds with the histological simplicity or complexity 
of the gland epithelium, and that this latter affords the only true 
basis for a classification of " red bodies." One of the most simple 
(though not the most primitive) conditions of the rete mirabile 
is when it is single and compact*. In this condition it may be 
separate from the glandular epithelium and ovoid in shape 
(Ophichthys iviberbis, fig. 9, e.g.), or contiguous with the glandular 
epithelium and circular in form (Gobius, fig. 22, e. g.), conical 
[Cepola, fig. 67, e. g.) or fan-shaped (Ophidium, fig. 48, e. g.), &c. 
In some cases the rete mirabile is divided into two, and here 
these are separate from the glandular epithelium and ovoid in 
form (Angitilla, text-fig. 53, j\Ji/ruSj Mztrcsna, Ophichthys serpens, 
(fee). In other cases the rete mirabile is divided up into several 
distinct par-ts, which often have a radiate arrangement : e. g. in 
Peristethas the rete is split up into some ten radiating strands 
(fig. 37 and text-fig. 57) and in some species of Gobius into seven 
or more. In most cases, however, the initial artery and vein 
supplying the bladder give off numerous arteries and veins in pairs 
(an artery and a vein to each pair, of course) at intervals so as to 
give rise either (a) to continuous sheets of rete mirabile, so to 
speak, very often two in number, one on each side, as in Corvina 
nigra (fig. 59), Sargus rondeletii (fig. 60), and Smaris vulgaris 
(fig. 45), or (6) to sheets divided up in variable degrees, as e.g. in 
Box boops, where the sheets are divided into outer and inner on 
each side (figs. 51, 52), or in Zeus faber, where there are three 
separate sheets — anterior, median, and posterior — on each side 
(fig. 62), or (c) to numerous separate small strands or tufts of rete 
mirabile, each strand supplying a small area of glandular 
epithelium : thus, in Co7'is jidis (figs. 56, 57) the rete is seen 
under the microscope to be subdivided into many small strands 
supplying the small area occupied by the gas gland ; in Trigla 
hirundo also a similar condition exists, but here, indeed, the twigs 
extend nearly over the whole interior of the bladder cavity (fig. 43) 
instead of being restricted to a small area ; in other cases where 
the tufts are larger and more separated, these can be seen with 
the naked eye, as, e. g., in Atherina hepsetus (fig. 53) and Perca 
(fio'. 69) and in the well-known case of the Cod (fig. 66), in which 
the subdivision of the rete mirabile reaches its maximum. 

Whether the condition of the rete mii^abile with which we 
started, i. e. as a single large well-defined body, separate from the 

* Reis (64) supposes that the horseshoe-shaped gland of Ophidium, e. g. (see my 
fig. 48 and compare it with figs. 59, 60, and 62), is the form of gas gland and rete 
from which all others are derivable, but it seems to me that the simple oval rete of 
Syngnathus in connection with an extensive area of glandiilar epithelium, is still 
more simple ; the most primitive condition of the epithelium and associated blood 
vessels is, of course, that of many freshwater teleosts in which practicallj' the whole 
of the internal unilaminar lining-epithelium of the bladder constitutes the gas gland, 
and the arterial and venous capillaries have not become associated to form a rete. 
In general it may be said that the more ditl'usethe rete and gland the more primitive 
their nature. 



GAS GLAXDS OF SOME TELEOSTEAN FISHES. 217 

glandular epitheliuai {Oi^hichthys miherbis, e. g.), is move primitive 
oi- more evolved than the condition which we found in the Cod, 
e. g., is a question I shall not discuss ; I will merely say that there 
are several reasons for accepting, with de Seabra (31), the latter 
view. The fact that Ophicht/njs and Anguilla possess a very simple 
type of glandular epithelium is no objection to this view, since 
the rete of Cejwla and other genera possessing complex types of 
epithelixnn is almost as simple. 

The glandular epithelium, as we have seen, may line the whole 
interior of the bladder (as in many Anguilliformes and freish- 
water teleosts) or be restricted to a small area of the internal 
surface {OpMdmm type). The Syngnathus type is more or less 
intermediate between these two conditions. 

The character of the glandular epithelium varies greatly. The 
cells composing it may be arranged to form [a) a single unfolded 
layer {Ab/rus, e. g.) ; (6) a layer thrown into simple or complex 
folds, so that channels are formed leading from the surface of the 
glandular epithelium to the bladder lumen {Anguilla, Gobiits, 
iSi/ngnathus, Trigla, Smaris); (c) a thick mass in which the deeper 
cells can only commvmicate with the bladder lumen by a system 
of large or small anastomosing channels (the intercellular lumina) 
which penetrate the cell-mass and open into the bladder cavity 
by small apertures situated on the internal surface of the 
glandular mass {Ophidium, Coris, e. g.). At present neither 
embrj^ology nor comparative anatomy provides evidence to justify 
the inference that this massive type of gland epithelium is in all 
cases derived from the folded condition — the folds on this view 
being supposed to have become elongated, laterally branched 
perhaps, closely apposed and fused at intervals to produce the 
cell-mass penetrated by narrow anastomosing channels. It would 
gratify me to be able to agree with Reis (64) who states that she 
is able to trace a complete developmental series of complications 
in the structure of the gas gland epithelium, the simple unfolded 
cell-layer forming the starting and the cell mass of Coris, Cejjola, 
or ^sovf the culminating points of the series; but, as I have already 
stated, the available embryological and anatomical evidence 
suggests that the two complex types of folded and massive gas 
gland epithelia were separately derived from the primitive layer 
of glandular cells by folding (i. e. the plane of cell-cleavage is 
always at right angles to the plane of the bladder wall) and bj'' 
thickening (the plane of cell- cleavage being parallel with the plane 
of the bladder wall) respectively. It is true that forms of gas 
gland epithelia occxu- apparently transitional in character between 
the folded and massive types [Trigla gurnardtcs, Smaris maiorii, 
6'. vulgaris), but I have supplied leasons for regarding these as 
glandular epithelia of the folded type which have become 
secondarily massive in form and so differing from the truly 
massive glands wliich never pass througli a folded stage in their 
development. Theie is no reason, however, why certain glands 
should not be truly intermediate in structure, i. e. in Avhich the 
cells during development divide simultaneously in planes both 



218 DE. W. N. F. AVOODLAKB ON THE 

perpendicular to and parallel with the plane of the bladder Avail, 
but I know of no examples of such. Subsequent embryological 
enquiry can alone prove which of these views is the correct one. 

The minute structvire of the cells of the gas gland epithelium is 
doubtless constant throughout the entire series of " red bodies." 
Gas bubbles are in all cases generated in the interior of the cells 
(usually in the neighbourhood of the nucleus) when the glands 
are active, and these bubbles are ejected by the ceils into the 
gland ducts and bladder cavity where they explode, the shattered 
walls of these bubbles giving rise to the masses of granidar matter 
already described. Vincent & Barnes analysed this granular 
matter (which, as before mentioned, must be caief ully distinguished 
from the disintegration products of the led blood corpuscles alone 
found in the blood-vessels) of the Cod and found it to consist 
principally of a nucleo-proteid ; and since the cytoplasm which 
forms the wall of the gas bubble is also composed for the greater 
part of this substance, the result of this single analysis is con- 
firmatory of the view just expressed, as is also the fact that a 
lai'ge amount of this granular matter is always associated with 
the presence of numerous gas bubbles. The iiitracellular ducts, 
which, with the other cytological features of the gas gland, have 
been so well figured by Bykowski & ISJusbaum (24) and Reis & 
Nusbaum (62) within the last few years, are doubtless also constant 
features of active teleost gas glands, though I am of opinion that 
some of these intracellular channels are due to the expulsion of 
bubbles from the cells and not purely nutritive as Reis & Nusbaum 
suppose. The intracellular capillaries, on the other hand, are 
permanent structures, and the fact that they occur so rarely as 
compared with the enormous number of these structures found in 
connection with the liver cells can only be explained by the 
relative inactivity of the teleost gas gland. 

I have already described and figured the hsemolytic disintegra- 
tion of the red blood corpuscles (erythrocytolysis as it may he 
called) which occurs in the blood-vessels associated with the 
active gas gland of Syngnathtis actts, and this same phenomenon 
is to be found in connection with other of my preparations of 
active gas glands and is doubtless a result of the gland's activity. 
In this process of haemolysis many of the erythrocytes become 
entirely broken up, first into lai-ge globules and finally into the 
granular matter which is present in such large quantity in the 
capillary lumina, whilst others are distorted in shape and only 
pa.rtially broken up, looking very much as if influenced by some 
toxic substance, as Jeeger suggests (see Part II.). This condition 
of the erythrocytes cannot be due to imperfect fixation since 
great care was exercised in the fixation and preservation of these 
gas glands ; moreover sections through other vertebrates — Elas- 
mobranchs e. g. — fixed by similar methods, exhibit the blood 
vessels as practically free from granular matter (the small amount 
sometimes present simply representing coagulated plasma) and 
the erythrocytes as retaining their form. 

Another fact which I wish to lay stress upon, especially in view 



GAS GLANDS OF SOME TELKOSTEAN FISHES. "219 

of the theory advanced in Part II., is the distribution of the 
granular matter in the capillaries of the rete mirabile. Careful 
examination of my sections* of Gohius niger, Gobius minutus, 
Syngnathus acus, Peristethus caiaphr actus, Gasierosteus splnachia, 
and some others, in all of which the gas glands are more or less 
active, had led me to the conclusions that granular matter is 
present in the venous capillaries of the rete to a far greater extent 
than in the arterial, and that the relatively small amount of 
granular matter present in the arterial capilLiries is chiefly 
situated next the distal pole of the rete — the pole next the gas 
gland. The small calibre of the arteries compared with the veins 
and the few series of sections of active glands, well fixed and well 
stained, wdiich I possess render it difficult to be absolutely certain 
of the above conclusions, but I am as certain as the limitations of 
my material will permit me to be. The bearing of these facts on 
the theory already outlined on page 198 is perhaps already 
apparent, but I shall reserve discussion for Part II. (see p. 232). 

As already implied, teleost gas glands aj-e best classified 
according to the character of the glandular epithelium. We 
divide gas glands into two groups: — (1) gas glands in which the 
glandular epithelium is a single layer, which may either remain 
unfolded or become folded in variable degrees ; and (2) gas glands 
in which the glandular epithelium is many-layered, the mass of 
cells being penetrated by fine ducts. With the exception of a few 
cases [Trigla gurnardus, Smarts maurii, S. vulgaris, and some 
others) in which the much-folded epithelium has secondarily 
assumed a massive character, this massive type of epithelium is 
probably not derivable from the folded type. Also in this second 
group of gas glands the rete mirabile is always contiguous with 
the glandular epithelium, reunion of the cajjillaries never 
occurring [Ophidium, Fierasfer, Box, Atherina, Coris, Balisies, 
Sargus, Caraiix, Charax, Herranus, Zeus, Gadus, Cepola, Esox, 
Perca, &c.). 

Group 1 is subdivisible into two minor groups : — 

(a) The rete mirabile of the gas gland consists of either one 
or two bodies distinctly separate from the glandular 
epithelium, the capillaries of the rete reuniting to form 
large arteries and veins before they come into con- 
nection with the glandular epithelium {Angtdlla, 
Myrus, Ophichthys, Muraena, &c.). 

(h) The rete mirabile of the gas gland is contiguous with the 
glandular epithelium, although a small amount of re- 
union of the capillaries occasionally occurs {Syngnathus, 
Gohius, Siphonostoma, Kerophis, Gasierosteus, Peri- 
stethus, Trigla, &c.). 

In concluding this account of the practical work connected with 

* Observations of tbe rete mirabile, when sectioned loiigitndinallj', are preferable 
from tins standpoint ; tbe arterioles are. of course, distinguishable from the venules 
bj thicker walls, by small calibre, and by the peculiar endothelium before mentioned 



220 DR. W. K. F. WOODLAND ON THE 

the present paper, I wish to express my great indebtedness to the 
Council of King's College, London, for generously allotting me 
the sum of thirty pounds out of the Government grant to the 
College for 1907 to defray a part of the expenses incurred by me 
during my visit to the Naples Zoological Station in April of that 
year. I wish also to express my indebtedness to the Naples 
Zoological Station Table Committee of the British Association for 
the occupancy of the table at Naples during the three weeks just 
referred to, and to the staff of the Naples Zoological Station for 
the large amount of trouble they took in supplying me with the 
material I required in connection with this and other subjects. 
Further, I wish to thank Professor A. Dendy, F.E..S., and 
Professor S. J. Hickson, F.Pt.S., for kind recommendations in 
connection with the above-mentioned grant and occupancy of the 
Naples table respectively. Professor J. P. Hill, D.Sc, and Dr. G. 
C. Chubb for recent friendly assistance, Mr. C. Tate Regan, M.A., 
for kindly advising me in connection with the systematic nomen- 
clature adopted in this paper (the nomenclature of Giinther's 
Catalogue), and Mr. C. Biddulph, who has considerably lightened 
the burden of preparing some hundreds of slides. Finally, I am 
also much indebted to the Council of the Royal Microscopical 
Society for kind permission to occupy their table at the Plymouth 
Marine Biological Laboratory during a week in August, 1909. 

Part II. — A few Suggestions concerning the Physiology 
OF the " Bed Bodies," 

It wasihy original intention to include in Part II. of this paper 
a resume of our present knowledge of the physiology of the gas 
bladder*, but although, with this end in view, I have become 
thoroughly acquainted with the whole of the vast literature 
dealing with the subject, yet other work prohibits me at present 
from thus devoting the time necessary for the composition of 
such a review. This being the case, I shall content myself with 
providing an outline sketch of the chief views which have hitherto 
been advanced to explain the mode of working of the "red bodies," 
in order that the reader may be in a position to estimate the 
value of the few additional suggestions I have to make. 

The several views which have been held concerning the origin 
of the three t principal gases — oxygen, nitrogen and carbon 
dioxide — contained in variable proportions in the gas-bladder 
cavity, can be classified into two categories : (1) the view (first 
attributed to Redi) that these gases are derived directly from the 
atmosphere, and (2) the views (lineal descendants of Needham's 
secretion theory %) that these gases are derived more or less 

* A recent account of the functions of the bladder will be found in Baglioni (18). 

t Argon is also stated to occur in the bladder. 

X It should be noted that authors in discussing the production of gas employ the 
term " secrete " in a verj' loose manner, some thereby meaning a true process of 
■ secretion, such as that which occurs in the sebaceous gland (Nusbanm & Eeis, 
e. ff.), others (Hiifner, Jaeger) merely meaning a process of pumping from the blood. 



GAS GLANDS OF SOMli XiiLEOSTEAN FISHES. 221 

directly fi'om the blood stre.im. The first view, abandoned by 
neai-ly all modern authors, is, however, persistently maintained 
by Tlulo(70, 71, 72). Thilo's principal contention, stated briefly, 
is that the blood of a fish is not suflicient in quantity to contain 
the amount of gas found in the bladder, and that even if this 
were the case the circulation of the fish is so feeble that the blood 
could not replenish an emptied bladder in the time experiment 
proves that it can be replenished. He therefore asserts that in 
all cases the bladder-gas must have been procured by the fish 
directly from the atmosphere : Physostomi can always renew their 
bladder -gas by rising to the surface of the water and passing air 
along the ductus pneumaticus, whilst Physoclisti are apparently 
under the hard necessity of absorbing a suificient supply when 
young, and therefore before the duct has degenerated, to last 
them throughout life, though Thilo further maintains that in 
many cases even adult Physoclisti are still able to procure fresh 
supplies b}^ passing air, either swallowed from the atmosphere or 
extracted in some inexplicable manner from the water, along the 
strand of tissue representing the vestigial duct. Thilo performed 
experiments and, according to his statements, obtained results 
which strongly support his view. He asserts, e, g. that he cut off 
the vascular supply of the previously-emptied bladder in Tinea (a 
physostome) and that in thirty hours it became refilled with gas. 
Seeing that the blood-supply was absent, he argixes that this fresh 
supply of gas must have been obtained from the atmosphere by 
passage thi-ough the pneumatic duct. I, however, cannot find 
that he analysed the gas produced under these conditions, and, 
since atmospheric air and secreted gas nearly always consist of 
oxygen, nitrogen and carbon dioxide associated in very different 
proportions in the two cases, his contention as to the soiirce of the 
gas receives very little real support from his expei-iments. A few 
of the many obvious objections to Thilo's hj^pothesis may be 
stated. In the first place, the percentage composition of the 
three gases contained in the bladder is, as just mentioned, quite 
diflerent from that present in the atmosphere, and in the cases 
of oxygen and nitrogen these gases are often present in such 
quantities as to exert a pressure many times greater than the pres- 
sures they exert in air — both of which elementary facts are fatal 
to Thilo's view. Thilo's contention that the blood of a fish with 
its feeble circulation is incapable of providing the quantity of gas 
required by the bladder of course begs the question at issue, and 
he might with equal force contend that the tissues of a siliceous 
sponge, for example, must be incompetent to deposit a massive 
siliceous skeleton, seeing that 100,000 parts of sea -water contain 
little more than one part of silica in solution and that diatoms 
a,nd other minute organisms are serious comj)etitors even for this 
small quantity (Sollas). Fiu^ther, Thilo's experimental results are 
not in accordance with those obtained by Hufner(38), Jaeger (45), 
and others, and, indeed, Thilo's own experiments failed to render 
his contention even pi'obable since the gland cells of the bladder 
would not necessarily cease their activity immediately a large 



222 DR. W. N. F. WOODLAND ON THE 

part of tlieir blood-supply was cut off. It is also certain that 
most physostomous fishes do not obtain their gases from the 
atmosphere vid the pneumatic duct, since the pneumatic duct is 
iiow known to be merely one of the two principal mechanisms 
(the other being the " oval ") employed by fish for the elimination 
and not the obtainment of gas. Finally, Thilo's theory fails to 
account both for the degeneration of the ductus pneumaticus in 
Physoclisti (since, according to Thilo, it is still functional) and 
for the presence of the complex "red bodies" which are the very 
structures needed by the fish on account of the feeble blood- 
circulation referred to by Thilo. 
.^ Theories which derive the bladder gases from the blood stream 
can again be grouped into two classes, viz. those which suppose 
that the blood gives up its gaseous constituents more or less 
directly to the bladder, the gas passing straight from the 
capillaries of the rete miiubile, or wall of the bladder, into the 
bladder lumen, and those which regard the gas gland as the 
special organ which extracts the gas from the blood. The first 
class of opinions, to some extent associated with the name of 
Moreau, is now quite out of date, though still to be found stated 
in some recent text-books. The second class of opinions comprises 
two quite distinct views as to the exact function of the gas gland 
- — two views which at the present time are held with equal tenacity 
by the schools represented by Jaeger and Nusbaum & Reis 
respectively. I shall first state briefly the view of Jaeger (44-48). 
Jaeger, following Hiifner (38), holds that the gas gland is 
pi-imarily a pumping apparatus, that is to say, an appai'atus for 
pumping the gases contained in the blood into the bladder cavity. 
The pressure exerted by the gases in the blood is, of course, 
considerably less than that exerted by the gases in the bladder *, 
and it is the function of the gas gland to force the gas from the 
blood into the bladder lumen against this superior pressure. 
Jaeger further supposes that the disintegration of a certain 
percentage of red blood corpuscles is effected by the secretion on 
the part of the gas gland cells of a toxin which is poured into the 
blood for this purpose. The object of so breaking up the ery- 
throcytes is to enable the gas gland cells, in some way not 
described, to lay hold of, with greater facility, the oxygen thus 
scattered in the corpuscle fragments. Indeed, Jaeger, following 
"Moreau, regards the gas gland as a mechanism essentially con- 
cerned with the pumping of oxygen — as an oxygen gland, in 
short, — a view confirmed both by the great development of this 
gland/ and by the disintegration of the er-ythrocytes in connection 
with bladders containing a large percentage of oxygen, and aJso 
by the contrary fact that in the bladders of Cyprinoids and many 
other freshwater fish which mostly contain nitrogen, the gas 
glands are absent — the ordinary squamous epithelial lining here 
being capable, without undergoing any special modification into a 

* JE. g. Jaeger (46) states that in deep-sea fish the partial pressure of the ox3-geii 
in the blood only amounts to about one-iifth of an atmosphere, whereas the oxygen 
in the bladder may possess a partial pressure of over fort}' atmospheres. 



GAS GLANDS OF SOME TELEOSTEAX FISHES. 223 

gland, of pumping in tlie nitrogen alone required. The reason for 
the special development of the oxygen gland in many deep-water 
fishes* is because oxygen alo»ie among the gases present in the 
bladder is capable (in virtue of its property of combining in large 
quantity with the haemoglobin of the blood) of being either rapidly 
produced (by the gas gland) or vapidly absorbed (l)y the " oval ' t), 
and this rapidity of production and absorption is essential in the 
case of fishes which undergo consi<lerable changes of pressui-e in 
the bladder. Deep-Avater marine or freshwater fish differ from 
most freshwater fish in that living in gi-eat depths of water those 
possessing migratory habits in a vertical direction require ap- 
paratus for adapting the volume of gas in the bladder to the violent 
changes of pressure expeiienced ; most freshwater fish, on the 
other hand, live in comparatively shallow water — in inland lakes 
and rivers — and thus lead placid lives, experiencing little or no 
changes of pressure in the bladder, and for these fish it is evident 
that thei-e is no need for the special development of an oxygen 
gland, which makes extrava.gant use of the gas which the fish 
requires for respiration : nitrogen and carbon dioxide sufiice t- 
"With reference to the spherical spaces pi'esent in the cells of the 
gas gland, Jaeger altogether denies that they represent gas 
bubbles : they are merely vacuoles such as are to be found in 
liver-cells, e. g. and have no connection with the production of 
gas. Jaeger himself, however, describes gas bubbles as being 
present in the ducts of the gas gland. Jaeger also refuses to 
admit that the large amount of granular matter found outside 
active glands in the bladder lumen is the product of the gas gland 
cells : in short, represents the broken-down w^alls of exploded gas 
bubbles. I shall discuss these features of Jaeger's theory later. 

Nusbaum & Reis (64, 55, 62-66), relying upon their extensive 
investigations of the cytology of the gas gland, formulate certain 
peculiar views § as to the exact mode of function of the gas gland. 
Jaeger, as already stated, holds with Hiifner that the gas gland 
is essentially a pumping apparatus, though he confesses entire 
ignorance as to exactly how the individual gas-producing cell 

* Only tbose deep-water fislies wliicli migrate vertically' and thns experience 
diflf'ereuees of pressure ; fish which remain at one depth, however great, obviously 
have no more need for " red bodies " than have surface fish (see p. 228). 

f Any sort of gas can, of course, be eliminated by tlie pneumatic duct. 

J Mile. C. M. L. Popta in her recent paper entitled " Etude sur la Vessie A^rienne 
dcsPoissons" (Ann. Sci. Nat. Zool.,t. xii, 1910, pp. 1-160) comes to the conclusion 
that " la vessie aerienne des poissons se developpe pour aider a la circulation de 
I'oxygene [et en general des gaz] dans le corps du poisson." I confess that this 
conclusion does not convey much meaning to me. I entirely fail to understand why 
the fish should develop an organ for the sole purpose of liberating oxygen into the 
blood in the same way I suppose that tlie liver liberates food material, and this is the 
only intelligible proposition which the author's conclusion conveys to me. Though 
the bladder undoubtedly in many cases performs this function incidentally, yet tlie 
fact that the bladder often contains other gases (nitrogen and carbon dioxide) the 
presence of which in the blood cannot be of the slightest advantage to the animal, 
shows that this storage function of the bladder cannot be its primarj' function. I 
cannot see that the author's conclusion in any way explains the diflcrdnt proportions 
in vrhich the three principal gases occur in the bladder. 

§ Unfortunately adopted in the last edition of Wiedersheim's ' Vergleichende 
Aiiatomie der "Wivbelthiere,' Siebentc Aufiage, 1909. - 



224 DR. W. N. F. WOODLAND ON THE 

extracts the gas from the blood and forces it against great 
pressure into the bladder : the supposition is, on his view, simply 
a legitimate deduction from the facts, and the details of the 
process, upon which he can throw no light, are of secondary 
importance. ISTusbaum & Reis, on the other hand, profess 
to have discovered in the cytology of the gas gland cell, some 
details of the pi'ocess which prove that the gas gland is not the 
mere pump which Jaeger assumes it to be. The view of ISTusbavim 
& Reis is briefly this : — Examination of the cells of the gas gland 
proves the existence of spherical spaces in the cytoplasm Avhich 
are not mere vacuoles as Jaeger assumed, but represent actual 
gas bubbles being formed in the cell. These gas biibbles are also 
to be found in the act of being ejected from the cells, and also 
lying freely in the gland ducts and bladder lumen, Jaeger himself 
necessarily agreeing that they are bubbles when found outside the 
cells, since liquid vacuoles could not possibly so pei'sist. Thus 
much is a matter of observation, but JSTusbaum & Reis contend, 
in explanation of these facts, that these bubbles of gas are pro- 
duced by the actual decomposition of the substance of the gas gland 
cells, just as gases are produced by putrefaction, and in proof of 
this cell-decomposition they describe the actual breaking-up of 
the cells — the nuclei * fraginenting in the manner described by 
Deineka for the Perch and the cytoplasm becoming converted 
into the masses of gi-anuiar matter found in the gland ducts and 
bladder lumen. The principal reason which leads xNusbaum &, 
Reis to this surprising conclusion is the supposed incompetence 
of the blood to supply the percentage of nitrogen gas found in the 
bladder. Among minor features of this hypothesis of Nusbaum 
& Reis, we may mention that they regard the disintegration of 
the red blood corpuscles both as a source of nutiition for the gas 
gland cells (the granular striping of the borders of gland cells in 
contact with capillaries and the intracellular channels penetrating 
the gland-cells being evidence of the cells absorbing this nvitrition) 
and as a source of the oxygen which, Avith the nutrition, 
is urgently required by the cells in connection with their rapid 
decomposition, 
f Thus the respective view^s of Jaeger and Nusbaum & Reis 

largely differ as regards the function attributed to the individual 
gas gland cell. Jaeger regards it as essentially a pumping 
mechanism ; Nusbaum &, Reis regard it as, primarily, mere 
substance to be decomposed into its gaseous and solid constituents, 
both being extruded into the bladder. 

I will now proceed to discuss these two views as a preliminary 
to a statement of my own views on the subject. Apart from 
omissions common to both theories, which I shall endeavour to 
lemedy shortly, the chief fault to be found with the theory of 
Jaeger is his refusal to recognize the origin of the gas bubbles in 
the cells of the gas gland. Concerning the existence of these 

* 1 maj' point out in this connection tliat during lisemolysis tlie nucleus of 
the red corpuscles appears to persist quite unchanged, even though the cytoplasm 
may become entirely disintegrated. 



GAS GLANDS O? SOME TELTi:OSTi;A^' FISHES. 



225 



structures tlier%,can be no doubt whatever (see Appendix B), 
They have been exhaustively studied by Bykowski and Nusbauni 
& Reis, and I have myself described and figured them in the 
present paper. Jaeger himself, as I have already stated, admits 
the presence of vacuoles in the gas gland cells, but, curiously 
enough, denies that they contain gas, and proi:iounces them 
similar in nature to the vacuoles in the cells of the liver. It is 
difficult to understand why this comparison should be made, since, 
from his own standpoint, it is the function of the gas gland cells 
to pump oat gas and not to store up glycogen or fat. But even 
adopting his suggestion that the supposed g-is bubbles are only 
vacuoles, we may remark that several authorities have described 
in the kidney " the formation of vesicles in the cells and appear- 
ances which indicate the discharge of these vesicles into the 
cavity of the tubules '' (37) ; and since kidney cells, like the gas 
gland cells, are supposed by many modern physiologists to derive 
most of their excreted substance by direct abstraction from the 
blood, comjjaratively few of the constituents of urine being 

Text-fig. 60 (x 1000). 




Vacuoles in the kidney cells of Nerophis, some apparently being expelled into 
the lumen of the tubule. 

manufactured in the cell, we may certainly conclude that even 
liquid vacuoles can be originated and discharged in a manner 
precisely comparable with that of the gas bubbles, and that if they 
contained gas instead of liquid they would be indistinguishable. 
I have figured some of the vacuoles present in the kidney cells of 
the fish Nerophis (text-fig. 60). It is evident that in histological 
preparations no sign of the liquid urine would be visible after 
expulsion from the cell, whereas the gas bubble with its walls of 
cytoplasmic material may, like a soap bubble in air, persist for 
some time before bursting. Consistently with his denial of the 
gaseous nature of the vacuoles in the gas cells, Jaeger was com- 
pelled to regard the gramilar masses lying external to the gas 
gland in the bladder lumen merely as a bye-product of the specific 
activity of the gas gland. 

If, however, we fully agree with Kusbaum & Reis as to the 



226 DR. W. N. F. WOODLAND ON THE 

origin of gas as bubbles in the cytoplasm of the gas gland cells, 
we as certainly disagree with the interpretation they put upon 
this phenomenon, viz. that the gas is produced as the result of 
the actual chemical decomposition of the cytoplasm. Like Jaeger 
we have found no evidence of that peculiar cell-disintegration 
which is supposed by Nusbaum & Reis to be the histological 
expression of the chemical decomposition of the cell. It is true, 
as Jaeger admits, that in the cells of active gas glands the 
cytoplasm (not the nucleus, which remains normal) often assumes 
a " hfird -worked "appearance — the cytoplasm looks "stringy" and 
contains numerous empty spaces — and considering that the 
process of continuous pumping of gas into the bladder must be 
exceedingly arduous, this is not surprising. It is also true, 
though not admitted by Jaeger, tbat the expulsion of bubbles 
from the cell into the bladder involves, to a considerable extent, 
waste of cell-substance. Each bubble, as already mentioned, 
possesses a wall of cytoplasm, and on the bursting of the bubble 
in the bladder lumen or gland duct, this wall, of course, breaks 
down and contributes to the mass of granular matter found in the 
bladder lumen and gland ducts. It must be confessed, therefore, 
that cell-disintegration occurs to a considerable extent and is 
associated with the production of bubbles of gas, but this 
mechanical disintegration of cell-substance is quite another thing 
from the hypothetical chemical decomposition of cell-substance 
postulated by Nusbaum & Reis (see Appendix B). Jaeger 
criticises this hypothesis of ISru.sbaum & Reis in a very effectual 
manner. He fii'st of all points out that it is impossible to regard 
the epithelium of the gas gland as analogous in its mode of 
working to a sebaceous gland, since the cells of the sebaceous 
gland decompose in order to produce a highly- complex substance 
chemically different from the substances supplied to them by the 
blood, whereas the cells of the gas gland give rise to the most 
simple of substances, viz. the gaseous elements oxygen and 
nitrogen which are supplied to them ready-made. In other 
words, the production of oxygen and nitrogen, unlike the secretion 
of the sebaceous gland, requires ho elaborate cell-metabolism, and 
there is therefore no reason for the gas gland cell-decomposition 
which Nusbaum & Reis affirm *. Further, according to Nusbaum 
& Reis, the blood corpuscles in breaking up supply the gas gland 
cells with oxygen for the decomposition of their substance, from 

* We may, indeed, comisave the process of gas-production with another process 
familiar to the present writer, in which a simple substance is also supplied to the 
cell and again liberated by that cell though in a difterent form, viz the deposition of 
calcareous spicules in various invertebrate groups. Calcareous spicules have, like 
gas bubbles, been regarded by some investigators as resulting from the actual trans- 
formation of cell-substance, but this view is not held by any modern zoologist. The 
spicule-secrcting cell is now regarded as a mechanism for abstracting the dissolved 
calcareous matter from the sea-water and of redepositing it, mixed with a variable 
minute amount of organic substance, in a crystalline form, the aggregate of calcite 
crystals constituting the spiciile. The work of the cell in this case is simply the 
abstraction and recrystallization of the dissolved calcareous salts — there is no question 
of cell-decomposition. The mode of working of the kidney cell may also be compared 
in this connection. 



GAS GLA^^DS OF SOME TBLEOSTEAN' FISHES. 227 

which decomposition oxygen is to be produced (!), but oxygen 
cannot be used up in decomposing tlie gland cells and yet be 
available to fill the bladder space. And as regards the supposed 
production of free nitrogen b}'^ the decomposition of the gas gland 
cells, Jaeger might have pointed out that such a fact would be 
unique in animal metabolism. Free nitrogen is not known to be 
liberated by aiiy katabolic process. The only source of the free 
nitrogen found in the bladder must be the small amount of that 
gas dissolved in the blood plasma. But Jaeger's chief objection 
to the theory of Nusbaum & lleis is the impossibility of accounting 
for the high pressure of the gas contained in the bladder, if this 
gas be simply produced by the decomposition of the cell-substance. 
The generation of carbon dioxide from chalk at once ceases, he 
points out, despite a high temperature, if the gas be allowed to 
accumulate in the calcination chamber, and in the same way the 
decomposition of cytoplasm would cease long before the liberated 
gas could exert a fraction of the jDressure usually found in the 
bladder *. Oppel (57), in a summary of papers by Reis & Nusbaum, 
also objects that whilst these authors strongly insist ujjon cell- 
degeneration yet they have not stated in what way the epithelium 
is regenerated after periods of activity, and I also have been 
unable to come to a conclusion as to the opinion held by Reis & 
Nusbaum on this subject. In one paper (62) these authors state 
in the last two sentences that they have observed mitotic figures 
in the gland cells, and conclude that regeneration of the gland is 
effected by ordinary cell-division ; in a subsequent paper (63), 
however, they appear to embrace the curious view that the 
amitosis first described by Deineka in Perca is a result of the 
violent cell-decomposition associated, in their opinion, with the 
activity of the gas gland, i. e. the nuclei, as well as the cytoplasm, 
undergo disruptive changes, and that this amitosis (55) leads to 
the production of new cells which take the place of those broken 
up! Needless to say, I fail to see any justification for this 
phcenix-like theory. My own researches have proved that the 
cells of the gas gland, when worn out like other cells of the body 
by a long period of activity, are replaced by ordinary mitotic 
divisions of the small cells at the base of the gland and that 
amitosis is only found in the later cell-generations. I may also 
mention in connection with this necrobiotic or decomiDosition 
theory of Nusbaum & Reis that, apart from other objections, it 
has always seemed to me to expect too much from the individual 
cell of the gas gland. The cell is not only requii^ed to commit 
suicide by undergoing decomposition but whilst undergoing 
decomjDosition it is to work energetically f in pumping the results 
of its own decay into the bladder cavity. Even the cell of the 

* This particular objection of Jaeger is invalid since the decomposition of cyto- 
plasm, being irreversible, would not be inhibited by pressure. 

t "The torniation of the gas bubbles in the cells is accompanied by fragmentation 

of the nucleus and simultaneous granular disintegration of the cell-plasma 

Condensation of the gas [in the cellj must accompany its formation since the lumen 
of the bladder is filled with gas at a considerable pressure." Translation of Reis (64). 



^ 



22S DU. \V. N. F. WOODLAXD OX THE 

sebaceous gland is not called upon for this degree of self-sacrifice. 
Finally, the researches of Deineka (29) on the intimate nerve- 
supply of the cells of the gas gland — ^it being stated that each 
gland cell is surrounded by a terminal cluster of nerve-filaments — 
and the known nervous control of the gas gland by two kinds of 
fibres running in the vagus and sympathetic nerves respectively 
(Moreau, 50, 51, 53 ; Bohr, 20 ; and others) render it very im- 
probable that such cells undergo the wasteful decomposition 
assumed by Nusbaum & Reis. 

I have now discussed the main features of the rival theories 
of Jaeger and Nusbaum & Reis. If we eliminate the errors 
and combine the truths of both with some observations of 
other authorities, we arrive at the following outline statement 
of the mode of working of the gas gland. The gas gland, in some 
unexplained manner, but probably, as Jaeger suggests, by the 
secretion of a toxin (which Reis. & JSTusbaum suggest he should 
have figured !) so influences the blood conveyed to it as to cause 
the disintegration of a certain proportion of red blood corpuscles. 
This fact of the disintegration is admitted by Bykowski and 
Nusbaum & Reis. The disintegration of the erythrocytes 
is primarily for the purpose of enabling the gas gland 
cells to "lay hold " of the relatively large quantity of oxj^gen 
contained in the blood. Oxygen, in consequence of its easy 
obtainment from the blood, is the all-important gas required by 
iishes which undei-go consideiuble and rapid changes of pressure 
in their bladder consequent on rapid changes in their vertical 
position, and the gas gland, considered as a special development 
of the lining epithelium of the bladder, is solely concerned with 
the rapid production of oxygen. The gas gland being essentially 
an oxygen gland is therefore most developed in those fishes which 
possess the greatest proportion of oxygen in their bladder gas ; in 
iishes like the Oyprinidie and some other freshwater families, on 
the other hand — iishes which exist in inland waters possessing 
little depth and which cannot, thei^efore, experience great changes 
of pressure in their bladder by vertical displacement, — nitrogen 
and carbon dioxide form the principal constituents of the bladder 
gas * and gas glands, in the ordinaiy sense of the word, and retia 
mirabilia are absent. It was formerly supposed that a large 
proportion of oxygen in the bladder was associated with the great 
depth at Avhich the fish existed — the greater the depth the greater 
the percentage of oxygen, — but apparently this is not altogether 
the case f. A large proportion of oxygen, as already stated, seems 
to be, for the most part, associated with the habits of those fish 
which frequently make considerable excursions in a vertical 
direction : if the fish sinks, the oxygen gland becomes active and 

* Cvprinus cai'pio contains about 94 per cent, of nitrogen in its bladder (Hiifner). 

t C'f. JSxocoetus volitans, tlie Flj'ing-tish, an essentially surface marine form, 
possessing, according to Humboldt (41), 94 per cent, of nitrogen in its bladder, with 
Coreqonus acronius, a fresliwater form living permanently at a depth of over 
seventy metres and possessing, according to Hiifnei", about the same percentage of 
nitrogen, and many other examples might be quoted. 



GAS GLANDS OF SOME TKLEOSTEAN FISUES. 229 

rapidly proLlu^es oxygen in considerable quantity in order to 
counteract the compression of the bladder due to the increased 
pressure and so to maintain the equality of the specific gravity of 
the fish with its medium ; if the hsh rises, either the absorption- 
organ known as the " oval " comes into action and returns the 
surplus oxygen in the swollen bladder to the blood, or, if the fish 
be a physostome, the pneumatic duct allows the surplus gas to 
escape to the exterior. Between the two extreme classes of 
bladder — the oxygen-filled bladder with " red bodies " and the 
nitrogen-filled bladder devoid of "red bodies" — there exist many 
transitional kinds containing a relatively small percentage of 
oxygen and feebly- developed "red bodies"*. As regards the 
exact mode of abstraction by the gas gland of the gases contained 
in the blood and their subsequent expulsion into the bladder, 
papers already published afford little or no information on the 
subject. In the case of those gases present in the bladder in 
minute quantities, it is probable that they leave the blood by 
simple diffusion: e.g. Traube-Mengai-ini (73, 74) and Priefer (60) 
showed that this was the case when hydrogen was dissolved in the 
water t. Ifc is evident, therefore, that the problem of gas- 
abstraction becomes of importance only when the partial pressure 
of any particular gas in the bladder exceeds that in the blood. 
That the gas gland cells act as a pump is certain J ; it is also 
certain that the gases first enter the cells in a dissolved condition 
and that, just as a scleroblast converts dissolved calcareous salts 
into a solid spicule,lso the gas gland cells cause these dissolved 
gases to appear in a gaseous form as bubbles in the cytoplasm §. 
In addition to this, however, the gland cells subject these gas 
bubbles to a considerable pressure, greater than that existing in 
the bladder, so that when the gas bubbles are expelled into the 
bladder lumen and experience a diminution of pressure they 
burst. As regards the varying composition of the bladder gas, it 
can only be concluded that the cells of the gas gland, like the cells 
of the kidney, exercise a selective power. That the oxygen 
pumped into the bladder is derived from the blood no one doubts, 
but Nusbaum & Reis, Thilo, and some others find a great 
difficulty in supposing that nitrogen (and carbon dioxide) is 

* E.g. Fercafluviatilis contains on an average 15 percent, of oxygen, 83 of nitrogen, 
and 2 of carbon dioxide (Hiifner) ; Lota vulgaris 65 per cent, oxygen, 30 nitrogen, 
5 carbon dioxide (Hiifner), &c., &c. For analyses of the bladder gases in many 
iish see the works of Biot (19), Contigliachi (27), Delaroche (30), Humboldt & 
Provencal (39,40), Hiifner (38), Richard (67) among others. The percentage 
composition of the bladder gas exhibits, as might be expected, considerable variation 
not only in different hidividuals of the same species of iish but in the same individual 
at different times. 

t It must be mentioned, however, that Humboldt & Proven9al (39) in 1809 per- 
formed this experiment of impregnating water with hydrogen and failed to detect a 
trace of hydrogen in the bladder, but probably their methods of analysis were too 
ciude. 

X In this connection Hiifner ventures to compare the gas gland cells with the 
contractile gland cells described by Drasch (32) in the neck-skin of the Frog, both 
being similarly supplied with nerves. 

§ Of. the gas vacuoles in Arcella and other Thecanioehida. 

Piioc. ZooL. Soc— 1911, No. XYI. 16 



230 DR. W. N. F. WOODLAND ON THE 

similarly derived. It is true that the amount of nitrogen 
dissolved in the blood is very minute as compared with the 
amount of oxygen, but, on the other hand, nitrogen does not, like 
oxygen, require to be produced at a rapid rate ; and since the 
supposition of Nusbaum & Reis, that free nitrogen is generated 
by the decomposition of cell-substance, is quite inadmissible for 
the reasons supplied above, the nitrogen dissolved in the blood 
can be the only source of that gas. It is well known that human 
blood, when suddenly released from gTeat pressure, develops 
bubbles of nitrogen owing to the inability of the blood to re- 
dissolve the gas immediately, and doubtless the cells of the gas 
gland also so act upon (though certainly not by a diminvition of 
pressure) the blood plasma bathing their substance as to comjoel 
the nitrogen to assiime a gaseous form. Once abstracted from, 
the blood stream, the nitrogen, in the case of those bladders 
containing nitrogen at high pressure, is pumped in the same 
way as, and in many cases with, the oxygen into the bladder 
lumen *. 

I now propose to oflfer some additional suggestions concerning 
the physiology of the gas gland which have occviri'ed to me 
during the course of my work. It is curious that not one of the 
investigators mentioned in the foregoing pages has attempted to 
explain in a satisfactory manner the striking conformation of the 
rete mirabile. A year ago I published in a short note (78) 
(based, as I have already explained, upon a mistaken conception 
of the teleost pancreas) a sketch of a new theory concerning 
the use of the rete which I have again outlined on a preceding 
page of this paper, but a,part from this I have met with no 
suggestion concerning the phj^siological significance of the rete 
mirabile bipolare geminum (or more simply rete mirabile duplex) t. 
I now propose to restate more fully this theory of mine, but 
before doing so I will mention the only previous hypothesis of 
which I am aware. Johannes Miiller (53) stated that in his 
opinion the retia mirabilia associated with the gas glands of 
teleost fishes (and Miiller was one of the first, if not the first, to 
distinguish the gas glands — " luftdriisen" — from the retia mira- 
bilia and to state their proper function as gas-producers) possessed 
the same utility as the various other kinds of retia mirabilia de- 
scribed by him, viz. to cause the blood stream to flow more slowly 
for some physiological purpose, and this opinion has been adopted, 
so far as I know, by all subsequent observers. Presumably 
the slowness of the blood stream in connection with the gas glands 
is supposed to be for the purpose of allowing the cells of the 
gas gland time in which to absti-act the gases present in the blood, 
and that this is one function of the rete- mirabile I myself do 

* Haldane (36) comes to the same conclusion. 

t The following suggestion concerning the use of the duplex rete associated with 
the gas glands may throw some light upon the phj'siology of the similar duplex 
retia mirabilia which are stated to occur in connection with other structures, 
e. cf. the " choroid gland " associated with the eye of many teleosts and the rete 
in connection with the liver of the Tunny. 



GAS GLANDS OF SOME TELdlOSTEAX FISHKS. 231 

not doubt.* But this explanation does not, in iny opinion, explain 
all the facts, since, if mere slowing down of the blood current 
is the one desideratum, then a rete solely connected with the 
bladder artery is all-sufticient. Further, it is evident that the 
explanation takes no account either of the fact that the bladder 
vein also forms a rete mirabile or of the still more remarkable fact 
that the retia on the bladder artery and vein respectively are both 
formed at exactly the same distance from the gas gland and their 
constituent arterioles and venules as intimately intermingled as any 
product of human manufacture purposely so designed could be. The 
suggestion Avhich I venture to ofter as to the use of this complex 
arterial and venous rete universally associated Avitli gas glands 
is the natural conclusion of the following considerations. The 
fact that some hundreds of the finest capillaries conveying 
blood to the gas gland are intimately intermingled with and 
closely apposed to a like number (another significant fact) of 
similar capillaries conveying blood from the gas gland is sug- 
gestive of the exchange between the two sets of capillaries of some 
substance necessarily of importance to the gas gland with which 
the rete is connected. This hypothetical substance we must 
assume both to be difi'used from the venous capillaries to the 
arterial (since diffusion in an opposite direction would not aJBfect 
the gas gland) and to be derived from the gas gland. The 
question which next arises is why this hypothetical substance 
should be poured into the arterial blood before it reaches the 
gas gland, and the only answer that suggests itself is that 
it is necessary for this substance to influence the arterial blood 
in some manner during its passage (made slower by the rete) 
from the rete to the gas glancl, so that by the time the arterial 
blood reaches its destination, its constitution has become altered. 
Now I have already stated that the gas gland, in the limited 
sense of the term, is essentially an oxygen-producing gland and 
that therefore the oxygen contained in the blood is the one 
element which the gas gland requires, from which fact we may 
conclude that the hypothetical substance referred to has some- 
thing to do with the giving up of oxygen by the blood to the 
gland. In the foregoing lines I have mentioned that such a 
hypothetical substance has already been postulated by Jaeger to 
explain the disintegration of the red blood corpuscles. We may 
therefore state as extremely probable suppositions that in the rete 
the venous capillaries contain a relatively large quantity of a 
toxin poui"ed into the blood by the cells of the gas gland, that 
this toxin diffuses from the venous capillaries into the arterial, 
and that whilst the artei-ial blood is slowly travelling from the 

* This function, however, cannot he exercised in the case of the retia mirabilia of 
the Eel, which are separated from the gas gland by a relatively few large vessels, 
since the reunion of the arterial capillaries on the side next the sras gland means of 
course the quickening of the blood stream. Neither can the Eel's rete mirabile 
possess the supposed function of the " carotid gland " of Amphibia, of " deadening " 
the heart pulse, since the gill capillary system of the fish must etfectively eliminate 
all trace of this ; on the other hand, the rete mirabile must certainly minimize the 
pressure of the blood supplying the gas gland, a result which, in the present instance, 
we must regard as a defect. 

16* 



232 DR. W. N. F. WOODLAND ON THE 

rete to the gland the toxin effects by a process of haemolysis the 
partial or total disintegration of a cei'tain proportion of erythi-o- 
cytes, the cytoplasm and oxyhjemoglobin pigment of each being 
scattered in the plasma, the former giving rise to the masses of 
granular mattei- so largely present in the blood returned from the 
bladder and the latter becoming dissolved in the plasma. 

Is there any evidence of this process actually taking place ? 
In reply to this question I am able to cite one fact, ah^eady stated 
on page 219 in Part I., which I think constitvites strong evidence in 
the affirmative. It is evident that if this process takes place 
then we may expect to find at the proximal pole of the rete, i. e. 
the pole remote from the gas gland, large quantities of granvilar 
matter (resulting from the erythrocytolysis above described) 
present in the venous capillaries but little or none in the arterial, 
and that towards the dit^tal pole of tiie rete (the toxin, on the 
hypothesis, having had time to diffi'use from the venous capillaries 
and influence the arterial blood) granular debris will also appear 
in the arteries. Careful examination of the rete mirabile asso- 
ciated with active glands has convinced me that this inferred 
distribution of the granular matter in the rete capillaries is a fact. 
Figure 32 (PI. lY.) represents a section taken across the proximal 
pole of the rete and shows the absence of the erythrocytolytic 
granules in the arteries ; a section across the distal end of the rete, 
on the other hand, shows that haemolysis of the arterial blood has 
commenced, erythrocytolytic granules being present. 

Oppel (56) puts a question which will be asked by many other 
critics of the foregoing theory of " red body " physiology. Why, 
he asks, is it at all necessary for the erythrocytes to be broken up 
in order that oxygen may be supplied to the cells of the gas gland ? 
In other words, if, as we know is the ciise, the blood readily gives 
up its oxygen to supply the ordinary tissues of the body, where 
is the necessity for the manufactvire of a special toxic substance 
in order to effect this same liberation of oxygen in the case of the 
gas gland ? I am not aware that Jaeger or any other author has 
given a definite answer to this question, but a consideration of 
the facts will soon supply one. In the first place, it is evident 
that the cells of the gas gland stand in a very different relation to 
the oxygen of the blood as compared with that of ordinary tissue 
cells, since whereas the latter are deficient in oxygen and require 
it solely for purposes of metabolism, the former are already 
saturated Avith oxygen aud only lay hold of it in order to con- 
centrate it and pump it into the bladder ; in other words, the 
cells of the gas gland require to get a very efliectvial "grip" on 
the oxygen which the tissue cells do not *. Secondly, realization 
of the conditions obtaining in the normal supply of oxygen by 
the blood to the tissues will enable us to understand the necessity 
for a toxin. The combination of oxygen with the haemoglobin 

* This necessity for the gas gland cells to be in the closest contact with the bicod 
is possibly accountable for the presence of the intracellular capillaries already 
describeil, perhaps also for the intracellular lumina as conveyors of exuded plasma 
into tlw celi-substance. The same phenomena are met with in the cells of the 
liver — an organ which also requires close contact with the blood—, the canaliculi 
vepvescntiiig the intracellular channels. 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 233 

pigment contained in the red corpuscles is, as is well known, a 
very loose one, and this loose combination is, according to modern 
views, only maintained by the maximum partial pi-essure exerted 
by the small amount of oxygen dissolved in the blood plasma. 
Furthei', it is important to notice in connection with the present 
subject that in the liberation of oxygen to the tissues of the body, 
it is the oxygen dissolved in the plasma which is immediately 
supplied to the tissues, and not the oxygen associated with the 
hajnioglobin. The oxygen of the oxyhamioglobin compovmd only 
becomes liberated as a consequence of the lowering of the partial 
pressure of the plasma oxygen caused by tissue a,bsorj)tion and 
when liberated meiely serves to replenish the plasma and is not 
conveyed directly to the tissues, so that the plasma with its 
limited solvent action on oxygen must be recognized as the all- 
important intermediary between the store of oxygen combined 
with the haemoglobin and the tissues. Now in the case of the 
cells of the gas gland, already employed in pumping oxygen into 
the bladder and certainly possessing all and moi'e than they 
require for metabolic purposes, it is difficult to imagine that they 
can absorb the oxygen dissolved in the pltisma in the manner 
employed by ordinary tissue cells. Even if they are so able, the 
small amount of oxygen so obtainable is quite insufficient for 
their purposes, since the oxygen, unlike the nitrogen and carbon 
dioxide, is required to be i^apidly produced in large quantities. 
The only alternative is, then, for the gas gland cells to seize upon 
the main source of the oxygen, viz. the oxyhsemoglobin, and so 
obtain in wholesale quantity what the plasma can only supply in 
retail*. This is effected, as we have seen, by the production of a 
toxin which, doubtless by a process of haemolysis, breaks up the 
red corpuscles into fragments and so liberates the contained 
oxyhsemoglobin into the plasma, the rete ensuring that this 
haemolysis and consequent solution of oxyhsemoglobin in the 
plasma is effected in time for the dissolved pigment to be available 

* This reduction of the blood to the primitive invertebrate condition in which the 
respiratory pigment is dissolved in the general plasma and not imprisoned in elastic 
discs (the erythrocytes) as in Vertebrates raises the question as to why, if the former 
condition enables the tissues to absorb the oxygen more readily, the latter condition 
has arisen. Apart from a few lamellil)ranch and other Mollusca and a few Poly- 
chajtes and PJioronis in which hasmatids have been described, all Invertebrates (and, 
according to Lankester, also Amphioxus and the Leptocephalus larva of the Eel, hut 
hajmatids have been stated to occur in the former and possibly exist in small number 
in the latter) carry the respiratory pigment, when this is present (absent in the 
tracheate. Arthropods, e. (C.), in the plasma, and it is difficult to understand what 
advantage accrues from preventing the dissolved oxyhiBmoglobin coming into direct 
contact with the tissues in vertebrate animals. It is of course possible that the 
indirect distribution of oxygen by way of solution in the plasma conduces to a more 
even and gradual supply to the tissues, especially in animals like Vertebrates in 
which most of tlie tissues are very remote from the limited respiratory area of the 
body surface ; in most Invertebrates, on the other hand, oxidation of the blood takes 
place over most of the body surface and the tissues are all practically simultaneously 
reached by the blood which, being contained in sinusoids rather than in capillaries, 
bathes them on all sides. The more rapid circulation of the blood and larger 
quantity of ha3moglobin in Vertebrates possibly compensate for the absence of 
h;emoglol)in in the plasma. As is well known, when hff'inoglobin is liberated into 
the plasma in Vertebrate blood, it is at once eliminated by the liver and kidneys. 
See Addenda (2). 



234 -DE. W. N. r. WOODLAND ON THE 

for absorption when the blood readies the gas gland. Is there 
any evidence that the cells of the gas gland absorb this oxy- 
haBmoglobin dissolved in the blood plasma ? It is not too much 
to say in reply that every preparation of a gas gland in at all 
an active condition does provide very substantial evidence as to 
the actual occurrence of this process. In every such preparation 
(see figs. 28-31, 50, 68, 73, e. g.) it is at once noticeable that the 
cytoplasm of the gas gland cells in contact with the capillaries — 
and usually the cells ai-e only separated from the blood by the 
thin endothelium — is of dense appearance and forms a perivascular 
zone quite distinct from the rest of the cell-protoplasm, a feature 
wdiich has been described and figured by all recent observers of 
gas gland structure — Jaeger, Bykowski, Nusbaum, Reis and the 
present writer (see Part I.). ISTusbaum & Reis hold that this 
appearance of the cytoplasm next the blood vessels is merely evi- 
dence of absorption from the blood of the nutritious matter afforded 
by the disintegration of the erythrocytes, but for the reasons just 
given we prefer to believe that the oxygen associated with the 
haemoglobin is the desideratum of the gland cells and that what- 
ever nutritive value ingested fragments of stroma may possess is 
quite a minor matter. However, the opinion of Nusbaum & 
Reis is of value in supporting our conclusion that the cells of the 
gas gland do actually absorb from the blood material liberated by 
the breaking-up of the erythrocytes. Examination of good 
preparations of active glands shows that this darkening of the 
cytoplasm of the individual gas gland cell situated next the blood- 
channel really possesses a striped appearance (PL IX. fig. 73) — 
Bykowski & Nusbaum, e. g., describe it as " die charakterische 
Streifung des Protoplasmas rings um die Blutgefiisse " — similar to 
that seen at the edges of the cells lining portions of the gut, in 
the Sertoli cells of the testis and in other cases, and this striping 
found in so many kinds of cells is proof of a process of absorption 
taking place. We do not suppose that the fragments of corpuscle 
substance composing the granular matter in the blood stream ai'e 
absorbed by the gas gland cells, but only the oxyhsemoglobin 
dissolved in the plasma, and the fact, which M^e have previously 
stated, that the veins of the rete mirabile are in active glands full 
of this granular matter is in accordance with this view. It can 
thus be proved by what practically amounts to actual demon- 
stration that the cells of the gas gland do absorb the dissolved 
oxyhEemoglobin directly from the blood (see Addenda (3)), and the 
natural inference is that this is employed for the supply of oxygen 
to the bladder. 

Judging from analogies provided by other classes of secreting 
cells, what possibly happens in the metabolism of the gas gland 
cell in the production of bubbles of ox3'gen gas from the absorbed 
oxyhsemoglobin dissolved in the blood plasma is that a,t that end or 
pole of the gas gland cell situated next the blood stream the 
dissolved oxyhpemoglobin forms a loose combination with the 
cytoplasm, this combination being merely a temporary linkage of 
the molecules of the two substances (similar, e. g., to the linkage of 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 235 

side-chains forming " anti-bodies," wliicli Ebrlich assumes to 
occur in his tlieory of immunity) and that later at the remote 
pole of the cell, i. e. the region of the cell remote from the blood 
strea^m, this loose temporary incorporation of the dissolved 
oxyhpemoglobin with the cytoplasm breaks down with the 
liberation of oxygen gas, which, as already seen, arises as bubbles 
usually in the vicinity of the nucleus. The slight decomposition 
of cell-substance which we may thus suppose to occur is evidently 
quite distinct from that decomposition postulated by Nusbaum 
& Reis. In our supposition the temporary linkage of the 
oxyhfemoglobin with the cytoplasm is merely to enable the 
cytoplasm to obtain a " grip " on the oxyhjemoglobin in order to 
dissociate the oxygen ; in the supposition of Nusbaum & Reis 
it is the cytoplasm itself which decomposes. 

A minor point remains to be mentioned. We have already 
stated that the veins returning the blood from the gas glanfl 
contain a relatively large amount of granular matter resulting 
from the h?emolysis of the red blood coi'puscles, and the question 
remains as to what becomes of this superfluous granular matter. 
It can only be said in rej)ly that probably a large portion of this 
disintegration material is eliminated from the blood by the liver, 
since we know that one function of the liver cells in all Verte- 
brates is to destroy degenerate corpuscles * and other waste 
material in the blood ; possibly also the spleen assists in this 
connection, though, judging from its histological appearance, I 
doubt it. I have also observed in several genera (A^erophis, 
Gohlus, Gasterosteiis, &c.) masses of cells lying to the outer sides 
of or between the kidneys in which a destruction of effete blood - 
corpuscles seems to be actively proceeding. It is, however, of 
little concern to us in what manner the blood is clarified in these 
teleostean fishes ; it suffices to say that the relative purity of the 
arterial blood in the bladder joroves that such elimination does 
take place. 

On re\iewing the foregoing pages it will be seen that the hypo- 
thesis concerning the physiology of the " red bodies " just elaborated 
has much to be said for it. In its essential features it is the theory 
of Jaeger and Ilufner, supplemented, however, and in some respects 
cori'ected, by the observations of Bykowski, Nusbaum, Reis, and 
other investigators, and slightly extended by the few suggestions 
made by the present writer. Its validity is assured by the number 
of diverse facts which it interprets, for not only does it explain 
the general fact of the existence of the " red body '"' as the only 
mechanism possible under the conditions for the rapid inflation 
of the bladder, but it also explains the minutest details of this 
mechanism, such as the perivascular striping of the component 
cells of the gas gland and the disintegration of the erythrocytes, 
and such a wide range of interpretation constitutes the criterion 
of a true theory, 

* These evytliroc.vtes in the liver have of course been largely dejirived of their 
oxygen by the gut tissues. 



236 DR. W. N. F. WOODLAND ON THE 

Appendix A. 

The Sources and Modes of Preparation of Material. 

A great part of my material was obtained by me during my occu- 
pation of the British Association Table at the Naples Zoological 
Station during April, 1907 ; additional specimens were sub- 
sequently sent to me from Naples and the remainder I obtained 
from the Plymouth Marine Biological Station. As regards 
methods of preparation, I may remark, first of all, that it is 
important to fix the lining-epithelium of the bladder and its 
special development, the gas gland, in a distended condition, and 
this is best effected by puncturing the bladder (in situ) posteriorly 
and immediately filling it with the fixative used, also pouring 
fixative on the outside of the bladder. By this means all shrinkage 
of the bladder wall is avoided on removal from the body. Another 
advisable precaution is to separate the lining-epithelium of the 
bladder, after it has been well fixed, from the outer layers of the 
bladder wall, since I found that in many cases the tougb muscle 
find connective tissue composing these latter cause difiiculty in 
obtaining thin sections across the gas gland. The fixatives which 
I employed chiefly were Zenker's Fluid, Corrosive Acetic, and 
Mann's Fluid (Distilled Water 100 c.c, Corros. Sub. 2-5 gm., Picric 
Acid 1 gm.. 40 % Formol 10-25 c.c. — a fixative which, in conjunc- 
tion with Borax Carmine and Picro-indigo-carmine, gave me most 
beautiful results). Some of my specimens were simply fixed in 
10 °/o Formol — a fixative which did not yield such good results as 
the others, though good enough for most purposes. I left objects 
in Zenker and Mann overnight, but in Cori'osive Acetic only for 
an hour or so. In many cases I fixed duplicate specimens in dif- 
ferent fixatives and was thus able to compare results. | Objects 
fixed with Zenker were well washed for several hours with Distilled 
Water and then, like the remaining objects fixed with other fix- 
atives, graded slowly up to 70 7o Alcohol (the pei-centages being 
in all cases made up with Absolute Alcohol and Distilled Water). 
Objects fixed with fluids containing Corrosive Sublimate were 
treated with Iodine at this stage. In most cases I subsequently 
stained in bulk with Grenacher's Borax Carmine, leaving the 
objects in for at least twelve hours, and toithout differentiation with 
Acid Alcohol, dehjAvKiQc]^ cleared, and embedded in Paraffin. The 
sections (both longitudinal and tra,nsverse in many cases), well 
stained with Borax Carmine, were then stained on the slide with 
Picro-indigo-carmine *. This stain is made up by adding one part 
of a satiirated solution of Picric Acid in 90 °/^ Alcohol (sat. sol. 
= circa 4'5 "/^ ) to two parts of a saturated solution of Grubler''s 
Indig-carmin in 70 7^ Alcohol (sat. sol. = circa 1 7^), and it is well 
to dilute this stain so obtained with twice its bulk of 70 7o Alcohol. 
The sections are placed in this stain for a short time, varying 
according to the thickness of the sections and depth of staining 
with Borax Carmine, from a few minutes to over half an hour. 

* I am iiidobted to Mr. A. D. Davbishire for recoinmendiiiff me this stain. 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 237 

The Picric Acid, of course, differentiates the.Borax Carmine, leaving 
scarlet nuclei, and the Picro-indigo-carmine stains the cytoplasm 
of the gas gland a dull green, the cytoplasm of red blood corpuscles 
and seci'etion products like zymogen granules a brilliant emerald- 
green, and connective tissue blue. Curiously enough, I did not 
always obtain these brilliant colour-contrasts with Zenker, espe- 
cially if the material had been preserved in Alcohol and Glycerine 
for some time after fixation, but fixation with Mann's Fluid always 
yielded the best results. Objects which did not stain well with 
the Borax Carmine I stained with Ehrlich's Htematoxylin and 
Picro-indigo-carmine, and obtained sufliciently good contrasts. The 
principal object to be attained by using Picro-indigo-carmine is to 
show up the blood corpuscles and granules. In exceptional cases 
I employed the Iron-hagmatoxylin method for nuclear details (in 
Pe7'ca, e.g.), also Kernschwarz for rendering visible cytoplasmic 
edges and hence the limits of intercellular and intracellular ducts 
(in Gadus, e. g.). As above mentioned, most of my gas glands were 
cut both transversely and longitudinally. 

Appendix B. 

On the Artificial Production of Gas Bubbles in Cells of the 
Gas Gland. 

Of the various methods available for compelling fish to produce 
gas in their bladder I have employed several, but only with one 
have I succeeded in detecting the gas bubbles in the act of being- 
formed by the cells. For the guidance of others I will first men- 
tion my unsuccessful experiments. I first experimented with 
Gobius pagaoiellus at Plymouth, subjecting this fish to increased 
pressure due to increased depth — one of the easiest methods of 
inducing gas-production. I took tank specimens from the Biolo- 
gical Laboratory on board the steam launch, and when anchored 
well out at sea I enclosed three or four specimens in each of five 
cages. One cage I let down to a depth of 30 feet and kept it 
there for one hour and a half ; a second cage 1 let down 60 feet 
for one hour, a thiid 90 feet for one hour, a fourth 180 feet for 
half an hour and a fifth 180 feet for two and a half hours. On 
examining sections of the gas glands of these fish I could detect 
no decisive differences in the gland cells. 1 am unable to explain 
why this experiment was a failure. Possibly even the maximum 
time allowed (2| hours) was too short in which to allow these cells 
to become active ; possibly also the fact that Gobius paganellus is 
a bottom form may have contributed to this negative result. 
Another experiment which I may mention was to stimulate the 
two vagus nerves of an Eel (Angioilla) with a battery for three 
hours, the two sympathetics having been cut and the bladder 
previously emptied and ligatured. At the end of the three hours 
the bladder certainly contained about one-sixth of its usual volume 
of gas, but I was unable to detect bubbles in the cytoplasm. I did 
not try the experiment of injecting fish with Pilocarpine nitrate 
or any similar drug. 



238 DR. W. N. F. WOODLAND OX THE 

The metliod by which I achieved success was that of attaching 
weights to some Perch in a tank in the manner described by Moreau *. 
The fish, if appropriately weighted, increase the volume of gas in 
their bladders in order to counteract the sinking action of the 
weights, but, according to my experience, it is all-important that 
the weights should not be too heavy, since if the fish are perma- 
nently attached to the bottom — if there is no hope of being able to 
rise — they apparently make no attempt at gas-production. The 
form of weight I employed was a half -hoop of lead weighing about 
5 gm. (appi'oximately one-tenth the weight of the fish, and this 
was probably too heavy) ; this I suspended round the lower half of 
the fish (by thin wiies tied dorsally) in the manner of a belt 
passing dorso-ventrally between the pectoral and pelvic fins, so 
leaving these perfectly free. After some twelve hours or moie 
the fish can swim about more easily and with a lighter weight it 
would regain all its usual mobility. If now the weight be removed 
the fish williise quite helplessly to the surface of the water, owing 
to the increased quantity of gas in the bladder. Unless the fish 
so rises after the experiment it is of little use to examine the gas 
gland. It is also, of course, necessary to compare the gas gland 
sections of the fish experimented on with similar sections of a 
control fish in the same tank. I weighted some six or seven Perch 
for 17 hours and at the end of that period the bladder of each was 
appreciably swollen, the fish having to swim vigorously in oi'der to 
keep below the surface after removal of the weight. I emptied 
the bladder in the usual manner by a small puncture posteriorly 
and in several cases refilled it with Zenker's Fluid ; in others I 
used a mixture of one volume of Glacial Acetic with tv/o volumes 
of Absolute Alcohol, and in others 1 "/^ Osmic Acid. Bladders 
fixed with Zenker I stained with Ehrlich and Picro-indigo-car- 
mine ; bladders fixed with the Glacial-Absolute mixture I stained 
with Iron-ha^matoxylin in order to study the details of nuclear 
structure ; bladders fixed with Osmic I stained with Ranvier's 
Picro-carmine. In all cases I made surface- view preparations, 
laying the bladder with its gas gland inside uppermost on the 
slide, in addition to cutting numerous sections of various thick- 
nesses of the Zenker and Glacial-Absolute material. The Zenker 
mateiial gave the best results from the gas bubble standpoint, 
probably owing to the absence of strong diflusion-currents during 
the process of fixation ; the Glacial- Absolute mixture, on the other 
hand, though perfect in its fixation of most parts of the cell, was 
doubtless somewhat drastic in its action on delicate structures like 
extruded gas bubbles. 

The sections of the Zenker material showed up the production 
of the gas bubbles in a very effective manner, the bubbles being 
clearly seen to arise in the cell-substance (not shown in text-fig. 61, 
p. 240) and to be extruded as foam-like masses all over the surface 
of the gas gland lining the bladder (PL IX. fig. 70 and text-fig. 61, 
the latter being a microphotograph). It must be remarked that 
only occasional cells were thus active, the majority not exhibiting 
* A more simple method, of course, is to empty the bladder with a trocai-. 



GAS GLAXDS OF SOME TELEOSTEAN FISHES. 239 

bubbles. Gas glands must be in a very active condition for the 
majority of cells to exhibit bubble- structures (see PI. V. fig. 35 of 
Gobius minuius). These bubbles were quite absent in the sections 
of non-weighted Perch used for the purpose of controlling the 
cxpeiiment. In my dacial- Absolute material, structures which 
are presumably bubljles (very unlike the bubbles sten in most 
preparations, however) are also to be found in the act of extru- 
sion, but these instead of being unmistakable foam-like masses 
ai-e bladder-like structures projecting from the cell-periphery 
(PI. IX. fig. 71 and text-fig. (32, the latter being a microphoto- 
grapli) and some are to be found lying free in the bladder-cavity. 
Possibly the action of the fixative is to cause all the small bubbles 
of the foam-like mass to run together to form one big bubble. 
In the surface-view preparations tixed by this method, large gas 
bubbles are occasionally to be found in the cytoplasm. All these 
bubble-structures are quite absent in the control material fixed 
with Glacial- Absolute. 

Apart from the existence of gas bubbles in preparations of the 
activated gland, another distinction from the non-active or dormant 
gland is the "worked" or "spent" appearance of many of the cells, 
a distinction w'hich I have tried to indicate in figures 71 and 72 
(71 represents cells of the active gland — cf. text-fig. 62 ; 72 repre- 
sents dormant cells). It v»'as this "spent" appearance, this 
attenuated and ragged condition of the cytoplasm, which suggested 
to Nusbaum & Reis that the cell substance actually decomposed 
into the bladder gases and granular masses, just as the cells of the 
sebaceous gland disintegrate to produce the secretion of that gland, 
and, indeed, at first sight of these exhausted cells of the gas gland 
the suggestion does not seem unreasonable. However, the fact 
that a cell which posseses but a small amount of protoplasm in the 
space bounded by its walls is not necessarily in a state of decom- 
position is shown by the majority of plant-cells, the cytoplasm 
of which is reduced to a " primordial utricle," by swollen adipose 
cells, by spicule-cells and by yolk-laden meroblastic eggs, to men- 
tion a few out of many possible examples, and since no amoiint of 
cytological technique can demonstrate the actual chemical decom- 
position Nusbaiim & Reis assume, this attenuation of the 
substance of the cells of the gas gland is not of very great 
significance, especially in view of the reasons for rejecting this 
suggestion of Nusbaum & Eeis alieady advanced in Part II. It 
must not be supposed that the cells of my activated glands alone 
wear this "spent" aspect, all the cells of the dormant gland 
resembling figure 72 ; on the contrary, the only difference between 
the cells of the two glands is one of degree, cells of the dormant 
gland being, on the whole, less ragged than those of the active. 
Only a gas gland completely at i"est (a condition probably rarely 
attained in a voracious fish like the Perch) would have all its cells 
in a resting condition. In most marine fish, however, which I 
have studied, nearly all the cells of the resting gland have been 
quite solid in appearance, only the cells of the active gland being 
attenuated. 



240 



DR. W. jST. F. woodland ON THE 

Text-fig. 61 (x circ. 1000). 




Microphotograph of transverse section across thin region of gns gland of Ferca (fixed 
with Zenker's Fluid), showing foam-like mass of bubbles (F.B.) on surface 
{cf. PI. IX. fig. 70). 

Text-fig. 62 (X circ. 1000). 




■*m-^ 



Microphotograph of transverse section across thick region of gas gland of Perca (fixed 
with Absolute and Glacial Acetic), showing extruded gas bubbles (B) on surface 
[cf. PI. IX. fig. 71). The " spent " condition of the cells is also shown. 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 241 

In connection with the two microphotogra.phs of gas bubbles 
hei'e reproduced as text-figures 61 and 62, I may say that these 
structures were in my preparations very difficult subjects for 
photogi'aphy, partly because of the inappropriate staining em- 
ployed and partly because bubbles shown in optical section of 
necessity bear but little resemblance to the real thing (this was 
especially the case with the foam-like mass indicated in text-fig. 61 ; 
cf. fig. 70). Nevei'theless, these microphotographs, taken for me by 
Mr. F.J. Pittock, of the Zoological Department, University College, 
are perhaps of value as affording impai'tial evidence of the 
appearance of an active gas gland epithelium, also of the " spent " 
condition of the cytoplasm. Microphotographs were also taken of 
intracellular bubbles, but, owing to the thickness and staining of 
the sections, were not suitable for reproduction. 

Addenda. 

(1) The statement that gas glands are normally quiescent may 
not be strictly nccurate ; it would perhaps be more correct to say 
that only on occasion do gas glands assume great activity. I 
make this remai-k in consequence of a communication from 
Lieutenant G. C. C. Damant, R.ISr., who kindly permits me to 
repeat it. From observations made under water during diving- 
operations and from the results obtained in catching fish on the 
hook at different states of the tide, Lieut. Damant concludes that 
Pout, e. g., remain fairly constantly at one level, viz., just off the 
bottom, whatever the state of the tide may be. If this be the 
case, it follows, as Lieut. Damant suggests, that the gas gland 
must become functional during each rise of the tide in order that 
a periodic increase of gas in the bladder may counteract the 
periodic increase of external pressure. 

(2) It seems probable, from suggestions kindly made to me by 
Dr. G. A. Buckmaster and Dr. G. 0. Mathison, that the principal 
factor in the evolution of erythrocytes in Vertebrates has been 
the necessity for a greatly increased quantity of hfemoglobin in 
Vertebrate blood as compared with Invertebrate — a necessity 
corresponding to greater functional activity. It is suggested that 
the amount of hfemoglobin required to be present in Vertebrate 
blood would be more than the plasma could possibly hold in 
solution, and that even if this were possible the viscosity of the 
plasma produced would seriously interfere with the other func- 
tions of the plasma, which, like the respiratory, have, in 
Vertebrates, increased in intensity. This lai^ge quantity of haemo- 
globin in Vertebrate blood being requisite and its presence in the 
plasma prohibited, the evolution of erythrocytes in which the 
hfemoglobin is imprisoned and combined with a proteid has 
apparently been the only alternative. As I have before remarked, 
the lack of direct contact between the tissues and the respiratory 
pigment in Vertebrates is compensated for both by the rapidity of 
the circulation and b)^ the amount of the respiratory pigment 
present, and doubtless also by the increase in amount of the waste 



242 DR. W. N. F. WOODLAND ON THE 

products of active tissues which act as reducing agents and so 
facilitate the abstraction of the oxygen dissolved in the plasma. 

• (3) I have adduced the striped darkening of the cytoplasm of 
the gas gland cells bordering the capillaries as evidence of the 
absorption of dissolved oxyhajmoglobin from the blood and that 
this is evidence of absorption is undoubted, but that it is absorp- 
tion of oxyhgemoglobin is of course only an inference from the 
hiemolycis of the blood corpuscles, the production of oxygen gas 
by the gland cells and other considerations. I wish to state 
here, previous to sending away the final proofs of this paper, that, 
judging from preliminary experiments with the Israel- Pappen- 
heim stain for hfemoglobin, kindly recommended to me by 
Dr. G. A. Buckmaster, I have been able to detect the actual 
presence of hsemoglobin in the cells of the active gas gland and 
its passage thereto from the blood stream, and Dr. Buckmaster 
has confirmed these preliminary observations of mine. I intend 
to repeat these observations on an extended scale on fresh 
material shortly, and until I have done so I do not wish the 
above statement to be considered as final. 

References to Pancreas Literature (pp. 198, 200 in text). 

1 Dale, H. H. — " On tlie Islets of Langerlians in the Pancreas." Phil. Trans. 
Roy. Soc. Lond., VoL cxcvii, 1905, pp. 25-46. 

2. DE Witt, L. M. — " The Morphology and Physiology of Areas of Langerhans in 

some Vertebrates." Jour. Exper. Med. New York, Vol. viii, 1906, 
pp. 193-239. 

3. DiAMAEE, V. — " Studii comparativi suUe isole di Langerhans de pancreas." 

Intern. Monatsschr. f. Anat. u. Physiol. Leipzig, Bd. xvi, 1899, 
pp. 165-209. 
3 a. Flint, M. — " Das Bindgewebe der Speicheldriisen nnd der Pankreas, etc." 
Arch. f. Anat. u. Entwick. Leipzig, Jahrg. 1903, pp. 61-106. 

4. Laguessb, E. — " Developpement du Pancreas cbez les Poissons Ossenx." 

Comptes reiidus hebdoni. des Seances et Memoires de la Soc. dc 
Biologie, Ser. 9, T. i, 1889, p. 341. 

5. " Pancreas intra-hepatique cliez les Poissons." Op. cit., Ser. 9, T. iii, 

1891, p. 145. 
6 " Sur la Formation des ilotsde Langerhans dans le Pancreas." Op. cit., 

Ser. 9, T. v, 1893, pp. 819, 820. 
rj " Developpement du Pancr(5as chez les Poissons Osseux." Journal 

de I'Anatomie et Physiologie, Paris, T. xxx, 1894, p. 79. 
g " Structure et Developpement du Pancreas d'apres les travaux recents." 

Ibid. pp. 591-608. 
g_ . " Sur le Pancreas du Crenilabi'e et particuli^rement sur le Pancreas 

intrahepatique." Kevue Biologique du Nord de la France, T. vii, 

1891-1895, pp. 343-360. 
\Q_ "Sur la Structure du Pancreas chez quelques Ophidiens et particuli- 

erement sur les tlots endocriues." Arch. d'Anat. Micr. Paris, T. iv, 

1901, pp. 157-218. 
11 Lane M. A. — " The Cytological Characters of the Areas of Langerhans." 

Amer. Journ. Anat., Vol. vii, Part 3, 1908, pp. 409-421. 
12. Leavaschew, S. — " Ueber eine eigentiimliche Verandernng der Pankreaszellen 

warmbliiliger Tiere bei starker Absonderungsthatigkeit der Driise." 

Arch. f. Mikr. Anat., Bd. xxvi, 1886, pp. 453-485. 
■ 13 Massaei. — " Sul pancreas di pesci." Eend. K. Accad. dei Lincei, Vol. vii, 

Fasc. 5, 1898, pp. 134-137. 
14. Opie, E. L. — " Disease of the Pancreas." (Lippincott) Philadelphia, 1903. 
1Q . " The Histology of the Islands of Langerhans." Johns Hopkins 

Bull., Baltimore, Vol. vi, p. 117. 

16 Oppel, a. — ' Lehrbuch der Vergleich. Mikr. Anat. Wirbeltiere.' Jena, Bd. iii, 

1900, pp. 800-818. 

17 Rennie J. — " The Epithelial Islets of the Pancreas in Teleostei." Quart. 

Jour. Micros. Science, Vol. xlviii, 1904-1905, p. 379. 



31. 



GA8 GLAXDS OF SOMi: TELEOSTEAN FISHES. 243 

Refereiiees to " lied Bodij " and oilier Literatare. 

18. B\&LioNi, S. — " Zur Pliysiolo2:ie der Sfliwimmblase der Fische." Zeitschr. f. 
Allg. Physiologie. (Voi-woni), Bd. viii, 1908, p. 1. 
V 19. BiOT.— r" Memoire sur la Nature de I'Air contenu dans la Vessie Natatoire des 
Poissons." Memoires do Physique et de Cliemie de la Soci(5t6 
d'Arcueil, T. i, 18U7, Paris. 

20. BoHEj C. — ''The Intiueiice of Suction of tlie Vagus Nerve on the Disengage- 

jnent of Gases in the Air-bladder of Pishes." Journal of Physiology, 
Vol. XV, 1894, p. 491. 

21. " Sur la Secretion de rOxygvne dans la Vessie Natatoire des Poissons." 

_ Conipt. rend. Acad. Sci. Paris, T. c\iv, 1892, p. 1560. 

22. Beidge, T. W. — " The Structure and Function of the Air-bladder in certain 

Fishes." Proc. Birmingham Phil. Soc, Vol. vii, 1889-1891, p. 144. 

23. Fishes, in ' The Cambridge Natural History,' Vol. vii, 1901. 

24. BxKOWSKi, L. & NusBAUM, J. — '■ Beitrage zur Morpliologie des parasitischen 

Knockentisches, Fierasfer Cuv." Bull. Acad. Sci. Cracovie, 1904, 
p. 409. 

25. CoGGi, A. — " Intorno ai corpi rossi della vescica natatovia di alcuni Telcostei." 

Mitt. Zool. Stat. Neapel, Bd. vii, 1886, p. 381. 

26. " Ueber den epithelialen Theil der sog. Blutdriisen in der Schwimm- 

hlase des Hechtes {Esox lucius)." Morph. Jahrb., Bd. xv, 1889, 
p. 555. 

27. CoNFiGLiAcm. — "Suir analisi dell' aria, contenuta nella vescica natatoria dei 

Pcsci." Pavia, 1809. 

28. COENING, H. K. — " Beitrage zur Kenntnis der Wundernetzbildungen in dem 

Schwimmblasen der Teleostier." Morph. Jahrb., Bd. xiv, 1888, p. 1. 

29. Deineka, D. — "Zur Frage ueber den Bau der Schvvimniblase." Zeit. wiss. 

Zool., Bd. Ixxvifi, 1904, p. 149. 

30. Delaeoche. — "Observations sur la vessie aerienne des Poissons." Ann. 
Mus. d'Hist. Nat., T. xiv, 1809, p. 184. 

DE Seabea, a. F. — "Sur les Corps Kouges des Teleosteens (Note pre- 
liminaire)." Bull. Mus. d'Hist. Nat. Paris, T. iii, 1897, p. 217. 

32. Deasch, O.— Archiv f. Anat. u. Phys., Physiol. Abth., 1889, p. 96. 

33. EiMEEY, C. — " Le Specie del Geiicre Fierasfer." Monog. 2, Fauna u. Flora 

des Golfes von Neapel, 1880. 

34. GouEiET, E.— "Da Role de la Vessie Natatoire." Ann. Sci. Nat. (5), 

Zoologie, T. vi, 1866. 

35. GuYENOT, E. — " Contribution a I'etude anatomique et pliysiologique de la 

Vessie Natatoire des Cypriuides." Compt. rend. Soc. Biol., T. Iviii, 
1905, p. 794. 
I 36. Haldane, J. S. — "Secretion and Absorption of Gas in the Swimming- 
Bladder and Lungs." Science Progress, Vol. vii (Vol. ii, n. s.) 1898, 
pp. 120 & 237. 

37. Howell, W. H.— ' A Text-book of Physiology.' Philadelphia, 1909. 

38. HiJFNEE, G. — " Zur phvsikalischen Chemie der Schwimmblasengase." Arch. 

f. Physiol., 1892, p. 54. 

39. Humboldt & Peoven^al. — Schweigger's Journal, 1809, Bd. 1, p. 119. 

40. Mem. delaSoc. d'Arcueil, T. ii, 1809, p. 400. 

4i. Humboldt. — ' Reise in die Aequiuoctialgegenden Deutsche Bearbeitung.' 
Stuttgart, Bd. 1, 1859. 

42. Jacobs, Che. — " Ueber die Scbwimmblase der Fische." Inaug.-Dissert 

Tubingen, 1898. 

43. — Tiibinger Zool. Arb., Bd. iii. 

44. Jaegee, a. — "Die Physiologic und Morphologic der Scbwimmblase der 

Fische." Pfluger's Archiv f. Physiol., Bd. xciv, 1903, p. 65. 

45. ■ — " Die Physiologie der Schwimmbkse der Fische." Biol. Centralbl., 

Bd. xxiv, 1904, p. 129. 

46. "Die Scbwimmblase der Fisclie." BericM der Senckenbergischen 

Naturforschenden Gesellschaft in Frankfurt-am-Main, 1904 (Oct. 
1903). 

47. • ■ " Zur Physiologie der Sch-wimmblase der Fische." Anat. Anzeig., 

Bd. xxix, 1906, p. 683. 

48. ■ " Erwiderung auf die in Heft 7/8 dieses Bandes des Anatoniischen 

Anzeigers erschiencne Entgegnung von Frau Reis und Herrn Nusbaum 
(Krakau) : 'Zur Physiologie der Scbwimmblase der Fische.'" Op. 
cit. Bd. XXX, 1907, p. 588. 

49. Milne-Edwaeds, H. — 'Le9onssur la Physiologie et I'Anatomie Compar^e.' 

T. ii, 1857, p. 368 et seq. 



244 DR. W. N. F. WOODLAND ON THE 

50. MoKEAU, A.—'"' Suv I'Air de la Vessie Natatoire des Poissons." Compt. vend. 

Seances Acad. 8ci., 1863. 

51. • " Reclievclies experunentales surles Fonctions de la Vessie Natatoire." 

Ann. Sci. Nat. (6), Zoologie, T. iv, 1876. 
52. ' Memoires de Physiologie.' Paris, 1877, T. xv, p. 494. 

53. MiJLLER, J. — " Ueber Nebenkiemsu und VVaiidenietze." Archiv f. Anat., 

Physiol, u. Med., 1840, p. 101. 

54. NusBAUM, J., & Re IS, K. — ■" Beitvage zur Anatomic und Physiologie des sog. 

Ovals in der Schwimmblase der Fische." Bull. Acad. Sci. Cracovie, 
Sci. Math, et Nat., 1905. 

55. NusBATJM, J. — " Zur Histologie der tatigcn Gasdviise und des Ovals bei den 

Teleostiern (eine Antvs'orfc an Alfred Jaeier)." Anat. Anzeiger, 
Bd. xxxi, 1907, pp. 169-174. 

56. Oppel, A. — Article on Atmungs-Apparat : Schwimmblase, in 'Ergebnisse 

der Anatomic und Entwickelungsgeschichte,' Bd. xiv, 1904, edited by 
Pr. Merkel und R. Bonnet, Wiesbaden. 

57. — Op. cif., Bd. XV, 1905. 

58. ■ — Op. cit., Bd. xvii, 1907. 

59. ■ — Article on Atmungs-Apparat : Schwimmblase, in the ' Lehrbuch der 

Vergleich. Mikr. Anat. Wirbeltiere.' Bd. vi, 1905, pp. 104-164. 

60. Peibfee. — ' Ueber die Funktion der Sclivirimmblase.' Berlin, 1892. 

61. QuEKETT, J. — "On a Peculiar Arrangement of Blood- Vessels in the Air- 

bladder of Fishes, vpith some Remarks on the Evidence which they 
afford of the True Function of that Organ." Trans. Micr. Soc. 
Lond., Vol. i, 1844, p. 99. 

62. Reis, K., & NusBAUM, J. — " Zur Histologie der Gasdriise in der Schwimm- 

blase der Knocken fische, zugleich ein Beitrag zur Trophospongieu- 
fvage." Anat. Auzeiger, Bd. xxvii, 1905, p. 129. 

63. " Weitere Studien zur Kenntnis des Baues und der Funktion der 

Gasdriise und des Ovals in der Schwimmblase der Knockenlische 
(Ophididffl, Percidas)." Op. cit., Bd. xxviii, 1906, p. 177. 

64. Reis, C. — " Weitere Beitrage zur Kenntnis der Gasdriise bei den Knocken- 

fischen." Bull. Internat. Acad. Sci. Cracovie, 1906, p. 771. 

65. " Materyaly do morfologii i fizyologi pecherza plawnego ryb 

kostnoskieletowych." Ivrali6w, Rozpr. Akad., 1906, pp. 639-670. 
(" Materialien zur Morphologic und Physiologie der Schwimmblase 
der Teleostier." Abhandl. d. Akad. d. Wissensch. Krakau, Bd. xlvi, 
^ 1907.) 

68. Reis, K. & Nusbaum, J. — " Erwiderung auf den von Dr. A. Jaeger in Bd. 29, 
No. 24 des Anatomischen Anzeigers veroff'entlichten Artikel : ' Zur 
Physiologie der Schwimmblase der Fische.' " Anat. Anzeiger, 
Bd. XXX, 1907, pp. 204-207. 

67. RiCHAED, J. — " Note suv le Gaz de la Vessie Natatoire des Poissons." Compt. 

rend. Acad. Sci. Paris, T. cxx, 1895, p. 745. 

68. Shore, T. W. — "Notes on the Origin of the Liver." Jour. Anat. Physiol., 

Vol. V, n. s., 1891. 

69. " On the Development of the Renal- Portals and Fate of the Posterior 

Cardinal Veins in the Frog." Op. cit.. Vol. xvi, u. s., 1901. 

70. Thilo, O. — "Die Entstehung der Schwimmblasen (Vorliiufige Mitteilung)." 

Biol. Centralbl., Bd. xxiii, 1903, p. 628. 

71. " Die Entstehung der Schwimmblasen." Korrespondenzbl. d. Naturf.- 

Ver. Riga, Bd. xlvii, 1904, p. 26. 

72. "Die Luftwege der Schwimmblasen." Zool. Anzeig., Bd. xxx, 1906, 

p. 591. 

73. Teaube-Mengaeini, M. — "Ricerche sui gas contenuti nella vescica natatoria 

dei Pesci." Nota 1 in ; Atti Accad. Lincei Rend., Vol. iii, 1887, 
2. sem. ; Nota 2 in : ibid., Vol. iv, 1. sem. ; Nota 3 in ; ibid. 

74. " Ueber die Gase in dev Schwimmblase der Fische." Arch. f. Physio- 

logie, Jahrg. 1889, p. 51 

75. Vincent, S., & Baenes, A. S. — " On the Structure of the Red Glands in the 

Swim-bladder of certain Fishes." Jour. Anat. Physiol., Vol. xxx, 
1898, p. 545. 

76. Woodland, W. — " A Suggestion concerning the Origin and Significance of 

the ' Renal-Portai System ' &c." Proc. Zool. Soc. Loud., 1906. 
77. • "The ' Renal- Portal System' and Kidney Secretion." Nature, 

June 13, 1907, pp. 151, 152. 
78. " On a New Gland in Tuleostean Fishes." Proc. Zool. Soc. Lend., 

1908, pp. 431-433. 



GAS GLAXDS OF SOME TELEOSTEAN FISHES. 245 

EXPLANATION OF PLATES II.-IX. 

[All figures of sections drawn with tlie aid of the camera lucida. In figures 4,9, 13, 
14, 15, 26, 38, 46, 49, 52 and 67, which, like most of the others, have been gi-eatly 
reduced in size during the process of reproduction, the capillaries of the rete 
mirabile have not been reduced to nearly the same extent as the other parts of 
the figures, since, had this been done, they would have been invisible.] 

AnguiJIa vulgaris (PI. II. figs. 1-8). 

Fig. 1 (X 18). Semi-diagrammatic transverse section through the anterior end of 

gas bladder (A. B.) and pneumatic duct (P.D.). G.E., glandular 

epithelium ot bladder ; D.E., epithelium of duct ; A., artery, and V., 

yein supplj'ing bladder. 

2 (X 18). Similar section, more posterior. The artery and vein have each 

divided, each pair of vessels consisting of an artery and a vein. 
3(X18). Still more posteriorly the ai-terj^ and vein of each pair have 
subdivided so as to form a mass of intermingled arteries and veins. 

4 (X 18). Each mass of arteries and veins has, just before the pneumatic duct 

joins the bladder, subdivided to form a bunch of minute parallel 
arterial and venous capillaries — the rete mirabile bipolare geminum 
(Muller) here seen in transverse section. 

5 X 18). Where the duct joins the bladder, the minute capillaries of the rete 

mirabile have reunited to a considerable extent, arteries with 
arteries and veins with veins, to form large vessels, which then 
recapillarize in order to suppli^ the gas gland. P.D., pneumatic 
duct which has now altered the character of its epithelium and 
become part of the bladder, P.B. 

6 (X 18). Semi-diagrammatic transverse section through the posterior region 

of the bladder, P.B. 

7 (X 500). Transverse section through the lining epithelium of the pneumatic 

duct. Note the large capillaries and squamous epithelium 
covering them. 

8 (X 500). Transverse section through the glandular epithelium of the bladder 

(gas gland). G.D., one of the gland ducts formed by the folding 
of the epithelium. 

Opliichthys {Sphcegehranclms) imberbis (PI. II. fig. 9). 

Fig. 9 (X cir. 10). Semi-diagrammatic transverse section through "red body," 
where the duct (P.D.) joins the bladder (B.). R.M., the single 
rete mirabile, the long axis of which is situated transversely in the 
ventral bladder wall. The capillaries of the rete are here seen in 
longitudinal section. 

Syngnatlms acus (PL II. figs. 10-13 ; PL III. figs. 14-21). 

Fig. 10 (X 27). Semi-diagrammatic transverse section through the anterior 

attachment of the bladder. A. and V., arteries and veins which 

subdivide to form the rete mirabile ; Ves., vessels not taking part 

in the formation of the rete ; B.G., masses of the diff'use pancreas. 

11 (X 27). The arteries and veins are here subdividing to form the rete mirabile. 

Figs. 12, 13 (X 27). The formation of the rete mirabile (R.M.). 

Fig. 14 (X 27). The anterior end of the bladder (A.B.) has just appeared. The 
i-ete mirabile (U.M.) is fully formed. 
15 ( X 27). The arterial and venous capillaries of the rete mirabile have reunite<l 
to form larger vessels which then recapillarize to supply the 
bladder epithelium (G.E.). B., bladder cavity. 

Figs. 16-18 (X 27). The vessels of the rete mirabile gradually disappear in supplying 
the bladder epithelium. 

Fig. 19 (X500). A fold of the "unthickened " bladder epithelium. B.L., gland 
duct between folds of epithelium ; G.S., granular matter present in 
only very minute quantity ; V.D., darkened zone of cytoplasm 
round capillarj- wall. 

20 (X cir. 1000). Transverse section through rete mirabile. Venous capillaries 

indistinguishable from arterial, S.G., disintegration-products of 
the erythrocytes. 

21 (X 1600). Stages in disintegration of the red blood corpuscles situated in the 

capillaries of the gas gland. S.G. as in fig. 20. 

Peoc. Zool. Soc— -1911, No. XVII. 17 



246 DR. W. N. F. WOODLAND ON THE 

Gobius niger (PI. III. fig. 22 ; PI. IV. figs. 23-32, 34; PI. V. fig. 33). 

Fig. 22 ( X 1). Appearance of the " red body " in the ventral bladder wall. 

23 (X 16). Semi-diagrammatic transverse section anterior to the "red bodj'." 

A. and V., the initial artery and vein which form the rete mirabile ; 
E.G., pancreas masses ; O.L., outermost layer of the bladder wall; 
G.E., glandular epithelium lining bladder cavity. It will be 
noticed that the vessels and pancreas lie outside the bladder wall. 

24 (X 16). More posterior section. The initial artery and vein have subdivided 

to a small extent. The outermost layer of the bladder wall has 
become broken through by the arteries, veins and masses of 
pancreas. 

25 (X 16). The arteries and veins, intermingled with masses of the pancreas, 

are rapidly subdividing to form the rete mirabile and the whole 
mass is now practically situated inside the bladder wall. 

26 (X 16). The rete mirabile (R.M.) has now commenced to supply the 

glandular epithelium (G.E.) and is situated wholly inside the 
bladder wall, i. e. internal to the outermost layer (O.L.). E.G., 
portion of the pancreas remaining external to the bladder. 

27 (X 330). Three folds of the bladder glandular epithelium in transverse 

section. C.R.M., minute capillaries of the rete mirabile; EU., 
gas-bubble; ICC, intracellular capillary. 

28 (X 800). Gas gland cell with intracellular capillary (ICC). Note the 

striped darkened zone of cytoplasm next the capillary. 

29 (X 800). Gas gland cells with inter- and intracellular capillaries (ICC). 

30 (X 800). Gas gland cell with intracellular lumen in transverse section. 

Intracellular ducts, when thus seen, are not always easy to 
distinguish from bubbles. 

31 (X 800). Gas gland cell with intercellular lumen (INT.CL.) in transverse 

section. 

32 (X 1000). Transverse section through anterior end of rete mirabile, showing 

the curious endothelium (END.) of the arteries in this region and 
the large amount of erythrocyte-disintegration granular matter 
contained in the veins. The arteries (A.) are here distinguishable 
from the veins (V.) by their smaller size, by their thicker walls, by 
possessing the curious endothelium and by being practically free 
from granular matter (S.G.). 

33 (X 800). A diict of the pancreas in longitudinal section. The numerous 

nuclei and the syncytial character of the wall are noticeable. A 
large amount of granular secretion-matter is present in the duct. 

34 (X 800). A duct of the pancreas in transverse section. 

Gobius minutus (PI. V. fig. 35). 

Fig. 35 (X 1000). Portion of the folded epithelium of the gas gland in a very active 
condition. Bubbles are seen being produced (usually near the 
nuclei) in the interiors of most of the cells, others are being 
extruded from the cells and one is shown Ij'ing freely in the 
bladder cavity. The large amount of granular matter lying in the 
bladder lumen external to the gas gland cells which has resulted 
from the bursting of extruded bubbles is noticeable. 

SipJioiiostoma typlile {rondeletii) (PI. V. fig. 36). 

Fig. 36 ( X 1000) . In addition to the erythroc3'tes there are shown three of the 
curious " white " corpuscles which are so striking in appearance in 
this fish, the cytoplasm being very dense. Tliese corpuscles are 
numerous. 

FeristetJms catapliractiis {Peristedion cataphractum) (PI. V'. figs. 37-41). 

Fig. 37 (X 1). Appearance of " red body" in ventral wall of bladder. 

38 (X 16). Semi-diagrammatic transverse section across the anterior end of the 

"red body" (see A-E, text-fig. 57, p. 202). S.E., squamous 

epithelium lining the greater part of the bladder; G.E., glandular 

epithelium; R.M., rete mirabile. 
Figs. 39-41 (X 1600). Cells of the gas gland showing formation of bubbles in 

cytoplasm. 



GAS GLANDS OF SOME TELEOSTEAN FISHES. 247 

Trigla hirimdo (PI. V. fig. 43 j PI. VI. figs. 42, 44). 

Fig. 42 (X 1). Lateral "red bodies" (L.R.M.) as seen in ventral wall of bladder 
(= lateral divisions of the elongated "red body," the median 
divisions being hidden bj' the muscle-bands, M.B., in the middle). 
G.E., the glandular epitlieliuni (too broad in figure). 

43 (X 16). Semi-diagrammatic transverse section through the middle of the 

"red body" represented in fig. 42. M.B., two strong muscle- 
bands on each side of the middle linej KM., rete mirabile ; 
L.R.M., lateral rete mirabile; G.E., glandular epithelium; 
S., strand of nerve-fibres supplying muscles. 

44 (X 250). Glandular epithelium of bladder in transverse section. B.L., gland 

duct. 

Smarts oulgaris (PI. V. fig. 46 ; PI. VI. fig. 45). 

Fig. 45 (X 1). Appearance of "red body " in ventral wall of bladder. 

46 (X 16). Semi-diagrammatic transverse section across anterior end of "red 
body." R.M., rete mirabile; G.E., gas gland epithelium; S.E., 
general squamous epithelium lining bladder; B.L., bladder cavity. 

Smaris maurii (PL VI. fig. 47). 

Fig. 47 (X 250). Pseudo-massive epithelium of gas gland in transverse section. As 
explained in the text, this type of epithelium is without doubt a 
derivative from the folded type and so differs from truly massive 
types of gas gland epithelia which are not secondarilyderived from 
a folded condition. The perivascular spaces seen in the figure are, 
in most cases at least, due to contraction. 

Ophidiam harbatum L. (PI. VI. figs. 48-50). 

Fig. 48 (X 1). Appearance of " red body " in ventral wall of bladder. 

49(X16). Semi-diagrammatic transverse section across "red bodJ^" S.E., 
squamous epithelium; G.E., gas gland epithelium; R.M., rete 
mirabile. Outer layers of the bladder wall stripped off. 
50 (X 800). Large gas gland cells with intracellular ducts (IC.D.), which 
possiblj^ represent the passages of bubbles to the exterior. B.L,, 
bladder cavity ; INT.C.D., intercellular duct. 

So.v hoops (PI. VI. fig. 51 ; PI. VII. fig. 52). 

Fig. 51 ( X 1). Appearance of " red body " in ventral wall of bladder. 

52 (X cir. 27). Semi-diagrammatic transverse section through the middle of the 
"red body" shown in preceding figure. R.M., rete mirabile; 
S.E., squamous epithelium; G.E.. gas gland epithelium; B.Ii., 
bladder cavity. 

Atlierina liepsetus (PI. VI. fig. 53; PL VII. figs. 54, 55). 

Fig. 53 (X 1). Appearance of "red body" in ventral wall of bladder (only the 
vessels and retia mirabilia are seen). 

54 (X 1000). The glandular epithelium in transverse section. 

55 (X 1000). The glandular epithelium of a young Atlierina (15 mm. long) 

showing incipient division of the cells in a plane at right angles 
to the bladder wall. C.T.C., connective tissue cells. Notice the 
large size of the nuclei as compared with those in the adult gland 
of another specimen. Though no capillaries are shown in the 
figure, yet thej' are plentiful in most parts of the gland. 

Coris jtilis (PL VI. fig. 56 ; PL VII. figs. 57, 58). 

Fig. 56 (X 1). Appearance of " red body " in ventral wall of bladder. 

57 (X 16). Semi-diagrammatic transverse section of "red bodi'." Lettering as 

before. 

58 (X 500). The glandular epithelium in transverse section. B.L., bladder 

cavity ; S.E., superficial squamous layer of the glandular mass ; 
BU., gas bubble; INT.C.D., narrow intercellular lumen. 

17* 



248 ON THE GAS GLANDS OF SOME TELEOSTEAN FISHES. 

Corvina nigra (PI. VII. fig. 59). 
Fig. 59 (X 1). Appearance of " red body " in ventral wall of bladder. 

Sarffws rondeletii (PI. VIII. fig. 60). 
Fig. 60 (X 1). Appearance of " red bodj^ " in ventral wall of bladder. 

JBalisfes capriscus (PI. VIII. fig. Gl). 

Fig. 61 (X 1). Appearance of " red body "in ventral wall of bladder. The "red 
body " is, in this case, situated posteriorly in the bladder, not 
anteriorly. 

Zeusfaher (PL VIII. figs. 62-64; PI. IX. fig. 65). 

Fig. 62 (X 1). Appearance of " red bodies " in ventral wall of bladder. R.M., rete 
niirabile ; G.E., gas gland ; M., anterior muscular band. 

63 (X 375). Transverse section across base of massive gas gland showing syn- 

cytial masses which probablj' represent the regenerating portion of 
tiie gland. Cell-outlines only become visible in the more periplieral 
regions. The syncytial nuclei divide by ordinary mitosis and are 
of several sizes, giant nuclei being among them. 

64 (X 666). Amitotic division of nuclei in some small giant cells, intermediate- 

sized cells and small cells of the gas gland. The nucleolus seems 
to be the first part of the nucleus to divide. 

65 (X 375). A binucleated giant cell surrounded bj^ the ordinarj"- small cells.. 

Comijare the magnification of this with the last figure. 

Gaclus morrJina (PI. VIII. fig. 66). 

Fig. 66 (X 27). Semi-diagrammatic transverse section across "red body." R.M., 
tufts of rete mirabile; G.E., gas gland epithelium; M.C., cap of 
connective tissue ; B.L., bladder cavit3^ 

Cepohi rnbescens (PI. VIII. fig. 67; PI. IX. fig. 68). 

Fig. 67 (X 1). Appearance of " red body " in ventral wall of bladder. 

68 (X 333). Glandular epitlielium in transverse section. The perivascular 
darkening of the cytoplasm is here very marked. The ducts of 
the gland are very thin and inconspicuous. 

Fercafluviafilis (PI. IX. figs. 69-72). 

Fig. 69 (X 1). Appearance of branched "red bod3' " in ventral wall of bladder, the 
portions of gas gland forming margins to the small fan-shaped 
retia mirabilia. 

70 (X 1000). A small portion of the gas gland of Perca (fixed with Zenker)- 

showing the evolution of bubble-masses from the cytoplasm in 
certain cells of the epithelium, consequent upon the experiment of 
weighting the fish (Appendix B, cf. text-fig. 61, p. 240). 

71 (X 1000). Cells of the gas gland of Per ra (fixed with the Glacial-Absolute 

mixture) exhausted by the activity of the gland {cf. text-fig. 62). 
One of tlie bladder-like structures is shown in connection ^^'ithone 
of the cells. 

72 (X 1000). Unexhausted, i. e. inactive cells of the gas gland of Perca (fixed 

by the same method) for comparison with those of fig. 71. 

Nerophis ceqiiorius (PL IX. fig. 73). 

Fig. 73 (X 666). Five gas gland cells of Neropliii in contact with a capillary. This 
figure, coloured to resemble my preparations as far as possible, 
shows in the gas gland cell the "striped zone of cytoplasm next the 
capillary wall (indicative of absorption) and the erythrocyte- 
disintegration granular material present in the blood. 



ox SKULLS OF OXEN FROM NEWSTEAD. 249 

12. On Skulls o£ Oxen from the Roman Military Station 
at Newstead, Melrose. By J. C. Ewart, M.D., F.R.S., 

F.Z.S.* 

[Received ami Road February 7, 1911.] 
(Text-figures 63-91.) 

Professor Marcellin Eoule, in his recent work on the Grotto of 
Grinialdit, states that Cuvier, Riitimeyer, Nehring, Gaudry, 
Boyd Dawkins, Duerst, and other naturalists who have studied 
the Quaternary Bovida; have regarded the Urus {Bos tauvus 
primigenius) as identical with our modern Bos taurus, of which it 
was probably the aircestor, and fi'om which it difltered only by its 
greater size. 

Though many naturalists since the days of Cuvier have 
dii-ected their attention to the history of Domestic Cattle, the last 
word has not yet been said about their origin, hence in dealing 
with the remains of cattle from the Roman Military Station 
at Newstead, Melrose, the investigator must still bear in mind 
that a fi.nal answer has not yet been given to the question — Are 
modern European cattle descended from the Urus, Bos taurus 
primigenius ? When discussing the origin of British cattle 
Pi'of . Hughes remarks : " Caesar mentions that thei-e were large 
herds of domesticated cattle in Britain, and we know from 
numerous excaA^ations into Roman and Roman-British rubbish- 
pits that these belonged not to the Urus but to Bos longifrons. 
This, then, is the native breed with which we must start in all our 
speculations as to the origin and development of British oxen. 
The Romans found that breed here and no other." 

Writing about the Celtic Shorthoi'n {Bos longifaons Owen, 
Bos hrachyceros Riitimeyer) Mr. Lydekker says, " It is, and can be, 
nothing but a variety of Bos taiirus " derived from the wild Urus 
-at a very remote epoch — ^" the occurrence of remains o'i an 
apparently similar breed in the prehistoric lake-dwellings of 
Switzerland suggests that the breed may have been established 
prior to the separation of Britain fi-om the Continent " % , 

■ Bos frontosus Nilsson, Lydekker also regards as a variety of 
the Urus, and as there was no other primitive Wild Ox in Europe, 
xind an Eastern derivation being in the highest degree improbable, 
Lydekker says that all the domesticated breeds of European 
cattle must trace their ultimate ancestry to Bos •j^'i'^nigenitos. 
While satisfied that the Domestic Cattle of Eui'ope are descended 
from Bos jjrhnigenitcs, Lydekker thinks it is quite probable that 
the origin of the humped cattle of India (Bos indicus) may be, at 
least in part, diflierent. 

* The author is indebted to Mr. James Curie, Priorwood, Melrose, tor the use of 
the Oxen skulls found at Newstead. 

f ' Les Grottes de Grimaldi,' tome i. fascicule iii., 1910. 
X ' Wild Oxen, Sheep, and Goats,' p. 18, 1898. 



250 



PROF, J. C. EWART ON 



Though Professor Fairfield Osborn thinks that the Domestic 
Ox, instead of being a direct descendant of Bos primigenius, is a 
descendant of Bos trochoceros of the Italian Pleistocene, he differs 




in no essential point from Lydekker, and, like Lydekker, believes 
British Shorthorn Cattle are descended from an indigenous 
Occidental race domesticated in Europe by the Neoliths*. A 
somewhat different view is taken by Prof. Boyd Dawkins, who 

* Osborn, ' The Age of Mammnls,' 1910. 



SKULLS OF OXEN FROM NEWSTEAU. 

Text-fiff. 64. 



251 




Front part of the skull of a Buffalo {Bos hubahis). 

The premaxilte {F.M.), as in the Anoa, extend upwards between the maxillae- 

and nasals. 

Text-fig. 65. 




Front part of skull of the Urus {Bos priinigeiiivs). 
In this skull the premaxillae are only slightlj' connected with the nasals, but in a 
skull in the British Museum the premaxillae extend nearly as far upwards 
between the maxillre and nasals as in the Buffalo (text-fig. 64) . 



252 



PROF. J. C. EWAKT ON 



says : " It is clear the domestic animals were not domesticated in 
Europe, bvit that they had already been under the care of Man 
probably for long ages in some other region. The Turf- Hog, the 
Celtic Shorthorn, the Sheep, and the Goat have been domesticated 
in the countries in which their wild ancestors were captured by 
the hunter in Central Asia. To this region also belong the 
Jackal, the Wild Boar, and the Wild Horse, and in ancient times 
the Urus. It is therefore probable that all these domestic animals 
came into Europe with their mastei's from the south-east — from 
the Central plateau of Asia — the ancient home of all the present 
European peoples." 

Text-fig. 66. 




Skull of a Urus in the Anatomical Museum, University of Edinbvirgli. 

The premaxilla has only a slight connection with the nasal, the forehead is flat, the 
temporal fossa is closed behind, and the horn-cores curve forwards at nearly 
a right angle to the forehead. 



Of Continental zoologists, Prof. Duerst has for some years been 
directing most attention to the origin of domestic animals. In 
his report on the Animal Remains found in Turkestan by the 



SKULLS OF OXEN FROM NEWSTEAB. 



253 



Pumpelly Expedition, Duevst says : " My recent studies on fossil 
remains of the bovines of the Indian Pleistocene have shown 

Text-fio'. 67. 




Front part of skull of American Bison. 

Tlie premaxillffi are short and far removed from the nasals. 

From a skull in the Anatomical Museum, University of Edinburgh. 



Text-fig. 68. 





P.M. 



Front and side views of the skull of a j'oung foital Ox. Nat. size. 
The prcmaxillae (P.J/.) reach the nasals {Na.), the froutals {Fr.) extend nearij' as 
far backwards as in the adult Anoa, and there is a large gap between the 
frontals {Fr.), parietals (Pa.), and supra-occipital (Oc). 



254 ON SKULLS OF OXEN FROM NEWSTEAD. 

me that the Indian (Narbada and Siwaliks) and China Taurina 
are the exact equivalents of the European Urus (Bos primigenms 
Bojanus), excepting some very slight variations pi'oduced by 
different geographical and local influences, so that the JBos nama- 
dicus Falconer and Cautley would represent the European Urus 
for the Asiatic Continent, especially the North Indian mountains 
and their neighbovxrhood" *. 

In recapitulating the results of his studies of the bovids of 
Anau, Duerst says: — " In the lower layers of period la from 
— 24 feet upwards there occur the remains of a wild Bos nmnadicus 
Falconer and Cautley. During period 16 there originates from 
this wild form a domesticated bovid, large and stately, provided 
with long horns. Judging from the measurements of the 
preserved bones this is absolutely the same Ox that was possessed 
by the Ancient Egyptians. 

" In the period II the size of the animal seems to have some- 
what diminished, unless possibly a smaller bovid reached Anau 
with the other newly imported domestic animals. It is, however, 
possible that this form of cattle of the culture II originated in 
a decline of the cattle-breeding of the later Anau-li ; as, indeed, 
the originally large long-horned Ox of the early Babylonians had 
already become small and shoi-t-horned in Assyrian times, and 
to-day, after a relatively shorter intei-val, shows a tendency to 
become hornless " f. 

Duerst goes on to say : — " The first remains of the long-horned 
breed [Bos taurus macroceros) belong at Anau about 8000 B.C. We 
find the same animal again about 3000 to 4000 B.C. in Babylonia 
and Egypt. At about 6000 B.C., however, we find that the large 
long-horned animal of Anau has become small and small-boned 
and has developed into a short-horned breed {Bos hrachyceros)X. 
Therefore all who do not believe in an autochthonous domesti- 
cation of the animals for each separate culture-sphere must admit 
that the original large and stately long-hornecl Ox of Anau was 
spread by tribal migrations before 6000 B.C. to Pei-sia and 
Mesopotamia and into Egypt and Central Africa on the one hand 
and on the other to India and Eastern Asia, where according 
to Chinese accounts it arrived in 3468 B.C. 

" Did the migration of the West occur only after the small 
breed had become established, i. e. about 6000 B.C., or even between 
6000 and 7000 B.C. w^hen the turbary sheep had formed ? To this 
question we have as yet no answer. We must, however, add that 
it was not in Anau alone that through unfavourable conditions 
of life the originally large and stately Ox was changed into the 
stunted and short-horned form {Bos taitrus hrachyceros). The 
same change took place in Mesopotamia, as one may easily per- 
ceive ni comparing the long-horned cattle of Chaldean or Sumero- 
Accadian times with the Assyrian small short-horned and the 

* 'Animal Remains from Excavations at Anau,' Carnegie Institution of Wash- 
ington, p. 361. 
t Duerst, op. cit. p. 369. 
J Sos hrachyceros is the same as JBos longifrons. 



Text-fiff. 69. 




• A. — Front view of a fa?tal Ox skull about the fifth month. The pvemaxillre {P.M.) 
reach the nasals (Na.), the frontals (Fr.) are long-, the panetals (Pa.) 
narrow, and the interparietals (Jp.) lie between the panetals and supra- 
occipital. Nat. size. 
."B. — Front part of same skull, to show relation of premaxilla? to nasals. Nat. size. 



256 PROF. J. C. EWART ON 

modern loose and sliort-horned or hornless cattle. There is, 
therefore, no reason for rejecting the assumption or hypothesis 
that the Ox of Anau, which about 7000 B.C. was undergoing 
this change of form, finally reached Central Europe, after its 
migration through Southern Russia and Eastern Europe, in the 
stunted form of Bos taurus hrachyceros" * i. e. in the small Celtic 
Shoi'thorn generally known in England as Bos longifrons. 

If the conclvisions ai-rived at by Duerst and others are justified, 
it follows (1) that all the modern domestic cattle — the humped 
breeds of India and Africa as well as the European breeds — are 
derived either from the Urus of Europe (^Bos taui'us 2}i'i'niigenius) 
or its reputed near relative the Urus of Asia {Bos namadicios), or 
are a blend of varieties or races of these two species ; and (2) that 
modern British breeds have been formed by crossing the Celtic 
.Shorthorn (Bos longifrons vel hrachyceros) — the small domesti- 
cated i-ace widely distributed over Britain in pre-Roman times — 
with Continental breeds (including shoi't-horned as well as long- 
horned varieties) introduced since the Roman invasion. Are these 
conclusions supported by the remains of cattle from the border- 
foi't occupied by Roman auxiliaries during the first and second 
centuries of the present ei'a ? 

Prof. Boule, in his recent work on the Grimaldi fossils, says the 
genus Bos (which includes the most specialized members of the 
Bovid family) seems to have been represented in Pliocene times 
by Bos planifrons and Bos acutifrons of the Siwalik deposits of 
India. Duei'st regards Bos planifrons t as the ancestor of both 
Bos priraigenius and Bos namadicus, but Riitimeyer thinks that 
though Bos p>lcinifrons may be ancestral to, it is only a variety of 
Bos primigenius, the European variety of Bos ncmnadicics. 

Though Bos p7'imigenius, like the Bison, only reached Europe 
in Quaternary times, it was soon widely distributed- — its 
remains occur in English Pleistocene deposits containing ElepJias 
antiquus and in deposits of a like age over the greater part of 
Eui'ope and also in North Africa. While Bos primigenms was 
extending its range over Eui'ope, Bos namadicus was spreading 
over Asia. 

It has hitherto been supposed that the Bison was more abun- 
dant in Europe during Pleistocene times than the Urus, bvit 
Prof. Boule believes that, at least in the vicinity of Grimaldi, the 
Urus was from the first as common as the Bison. 

About the colour of the Urus nothing absolutely certain is 
known, but from drawings of Mediaeval, as well as Palaeolithic 
ai'tists we can form a fairly accurate conception of its conformation. 
A picture, believed to have been made in Bavaria about 1500 a.d,, 
probably brings out the chief points of Bos 2>'>^i')nigeniibs %. 

* Duerst, op. cit. p. 440. 

t According to Duerst's latest view there is no real difference between Bos 'plant- 
fi'ons of Riitimej^er and JBos acutifrons of Lydekker, but at one time he believed 
Bos acutifrons was the predecessor of Bos namadicus, to which the Bibovine (Gaur 
and Banting) group and especially the Indian Zebu were related. 

J This picture is reproduced in the ' Oyclopffidia of American Agriculture,' vol. iii. 
1900 ; the Urus apparently survived in Poland up to 1627. 



SKULLS OF OXEX FROM NEWSTEAD. 

Text-%. 70. 



257 





A. — rront view of Calf's skull at birth. 
The premaxillaj {P.M.) fail to reach the nasals {Na.). The parietals and inter- 
parietals have coalesced to form a narrow plate {Fa.) between the frontals 
{Fr.) and supra-occipital. 

B. — Front part of same skull. 

The premaxilla {P.M.) is short and some distance from the nasal (iVa.). 

Text-figs. 69 & 70, from specimens belonging to Prof. Charnock Bradley, D.Sc. 



258 



PROF. J. C. EWART ON 



The Urus was widely distributed in Britain in Neolithic times, 
but the examination of Roman and Roman-British stations has 
hitherto afforded no evidence that Bos pri7nigenius still survived 
in England at the Roman invasion. 

A number of more or less complete skulls of the Urus have 
been found in Scotland, and fragments of skulls, limb-bones, or 
horn-cores believed to belong to the Urus have been found 
in almost every county in Scotland between the Solway and the 
Pentland Firths, and some horn-cores found in Orkney are so 
large that it is assumed they belong to Bos jJ^nigenius. 

Text-fig. 71. 




Front part of skull of a polled Aberdeen-Angus Ox, with premaxillffi {P.3£.) extending 
nearly as far up between tbe maxilla {Mx.) and nasals (Na.) as in the Buffalo 
(text-fig. 64, p. 251). From a specimen in the Royal Scottish Museum. 



Though there is no evidence that the Urus survived long 
enough in England to give rise to the Chillingham and other 
" wild" white park cattle, it has been suggested that a sufficient 
number survived in the Caledonian forests to found the Oadzow, 
Atholl, or other Scottish herds of " wild " cattle. The bones from 
Newstead, however, afford no evidence that the Urus still survived 
in Scotland when the Romans constructed the border-fort during 
the later part of the first century a.d. 

The skull of Bos taurus jjrimigenms is in some respects more 
highly specialized than that of any other member of the Bovidse. 
Hitherto in studying bovine skulls a A^ery considerable amount of 
attention has visually been directed to the position, size, and 



SKULLS OP OXEN FROM NEWSTEAD. 



259 



direction of the horns. I shall, however, especially refer to the 
premaxillse, occiput, and tempoi'al fossae. 

1. The Preinaxillce. — In some Zebras (e. g. the true Burchell 
Zebra) the premaxilla may only be connected with the nasal for 
a distance of 12 to 15 mm., while in others (e. g. a variety which 
lives near Lake Baringo) the connection between the nasal a,nd 
the premaxilla may exceed 50 mm. Differences in the food may 
accovmt for the premaxillae being long in some cases and short in 
others. The premaxillae may i-equire to be firmly wedged in 
between the maxillte and nasals in varieties in the habit of feeding 
during part of the year on coarse hard food, but only slightly 
connected with the nasals in varieties which usually consume soft 
green herbage. 

Text-fiff. 72. 




Skull of a Syrian Ox with vestigial horn-coves and a forehead like that of flat-polled 
Aberdeen- Angus cattle ; the premaxillaa (P.M.) as in text-fig. 71 extend far 
up between the maxillaa {3Lv.) and nasals (Na.). From Wilckens, 'Natur- 
geschichte der Haustiere.' 



In the Anoa (Bos dejoressicornis) of Celebes and in the wild 
Indian Bufialo {Bos bubahcs) the premaxillae are firmly wedged in 
between the maxillae and nasals. In the Anoa (text-fig. 63, p. 250) 
the total length of the premaxilla is 1 05 mm., and its connection with 
the nasal is 35 mm. ; in the Bufl:alo (text-fig. 64) the total length 
of the premaxilla may be 173 mm., and its connection with the 
nasal 40 mm. 

This long intrusion of the premaxilla between the nasal and 



260 PROF, J. C. EWART ON 

maxillary bones doubtless increases the strength of the front 
part of the jaw which supports the horny pad against which the 
lower incisors bite. As the Anoa is partial to the neighbourhood 
of water, it probably feeds on coarse grasses like its ally the 
Tamarau of the island of Mindoro, which is said to browse on 
sugar-cane. 

The Indian Buffalo lives in the neighboui-hood of swamps and 
jungles, and probably also feeds on reeds and coarse grasses — • 
food which necessitates long, firmly secured premaxillse. 

In the Catalogue of the Ungulata in the British Museum 
Dr. Gray says that in the tiaie Oxen "the pi-emaxillee are large 
and always extend upwards into the triangular space between the 
maxillge and the nasals and consequently articulate with both 
these bones as in Buhalus ; in the genus Bihos, on the other hand, 
the premaxillse are small and are attached only to the distal 
extremity of the maxilla and are separated by a considerable 
interval from the nasals." As will appear below, the premaxillse, 
instead of being always long enough in true Oxen to extend 
upwards between the maxillfe and nasals as in the Buffalo, are 
sometimes so short that they fail to reach the nasals as in the 
Bison (text-fig. 67, p. 253). 

In Bos 2^rimigenius the premaxillse, in all the skulls I have 
examined, reach the nasals. In some cases the connection with 
the nasals is only 5 mm., but in a Urus skull in the British 
Museiim the premaxillee extend nearly as far upwards between 
the nasals and the maxillae as in the Buffalo (text-fig. 64). The 
extent of the connection between the premaxilla and the nasal 
in a Urus skull in the Anatomical Museiim of the Univei-sity of 
Edinburgh is shown in text-figs. 65 & 66. 

The premaxilla is shorter and further removed from the nasal 
in the Bison than in aiiy other member of the Bos genus (text- 
fig. 67). According to Major Heber Percy, the European Bison 
" are fond of grazing on a coarse aromatic kind of grass known as 
Zubr gi^ass." Others state that " they are equally fond of browsing 
on the leaves, young shoots, bark, and twigs of trees," and that 
" in winter they are driven to subsist entirely on buds, twigs, bark, 
and such patches of dry grass and fern as remain " *. 

The prairie Bison of America apparently fed chiefly on grass, 
hence doubtless the necessity for the extensive migrations, but the 
American woodland Bison seem to " subsist chiefly on the leaves 
and twigs of the birch and willow " t. It is conceivable that 
owing to the softer nature of the food of the Bison the necessity of 
having the premaxillse firmly wedged in between the nasal and 
maxillary bones no longer exists. 

In a very young Domestic Ox skull (breed unknown) the pre- 
maxilla (text-fig. 68) bears the same relation to the nasal as in 
the Urus represented in text-fig. 66 ; in an older skull it occupies 

* Lj'deklver, ' Wild Oxen, Sheep, and Cattle,' p. 77. 
f Lj'dekker, op. cit. p. 91. 



SKULLS OF OXEN' FROM XEWSTEAU. 



261 



the angle between the maxilla and tlie nasal (text-fig. 69, A, B) ; 
but in a skull believed to be tliat of a new-born calf there is 
a considerable gap between the preaiaxilla and the nasal (text- 
fig. 70, A, B). In some of the skulls from the JSTewstead Fort the 
premaxilUe are large and have nearly as extensive a connection 
with the nasals as in the Buffalo ; in others they are short and 
separated from the nasals by a considerable interval. 



Text-li 




Front part of skull of polled XewsteaJ Ox in wliicli the upper end of the 
prcmaxillaj (P. 31.) bifurcates. 

In a Newstead polled skull the premaxillse are long and more 
extensively connected with the nasals than in tlie Urus skull 
shown in text-fig. <56. In some Aberdeen-Angus cattle the pre- 
maxillse are short, in others they are so long and so extensively 
connected with the nasals that they reach to within 25 mm. of 
the lachrymal bones (text-fig. 71); in a Buffalo with premaxillfe 
of the same length (text-fig. 64) the lachrymals ai'e 75 mm. from 
the premaxillfe. Apparently in some Syrian cattle the premaxilke 
extend well up between the maxilke and nasals. In a skull of a 
Syrian Ox figured by Duerst* the premaxillse henv the same 
relation to the nasals and lachrymals as in the Angus skull 
figui-ed. As this Syrian Ox h;id small imperfect horns (text- 
fig. 72), and as the skull apart from the horn-cores resembles 
the skull of the flat-polled Angus strain, it may represent a phase 
in the evolution of the Aberdeen -Angus breed. The larae 



* Wilckons 

Proc. Zool. Soo. 



■ Naturji-i'schiclite der Haustiere,' p. 311, lODo. 

-1911, No. XVIII. 18 



262 



PEOF. J. C. EWART OX 



prem<axillfie, coiii3led with the fact that the parietal extends 
well on to the forehead, suggest that Aberdeen-Angus cattle 
have in part sprung from a race domesticated in Central Asia. 
While in two of the skulls of black polled Aberdeenshire cattle 
hitherto examined the pi-emaxillse have an extensive connection 
with the nasals, in the skull of a white polled " wild " Cadzow Ox 
as in a black polled Galloway, the premaxillre bear practically the 
same relation to the nasals and the lachrymals as in Bos 2^rivii- 
genius, represented in text-fig. 65. 

Text-fi2-. 74. 




Front part of a horned Newstead skull in wliicli the premaxilte {F.M.) are short 
and fail to reach tlie nasals {Na.). 



In most of the Newstead skidls examined the premaxill^e are 
absent, but when the maxillse are present it is usually possible to 
say whether or not the premaxillse had reached, or all but reached, 
the nasals. The}^ probably reached the nasals in at least 90 per 
cent, of the skulls of the Celtic Shorthorn {Bos longifrons) tyjDe, 
and in about 70 per cent, of the skulls belonging to long-horn and 
cross-bred animals. In the Celtic Shorthorn skulls the premaxilla^. 
though long and almost in contact with the nasals, instead of 
ending in a wedge-shaped process, are bifurcated and send a 
process backwards over the maxilla as well as one upwards between 
the maxilla and the nasal (text-fig. 73). In having the proximal 
end forked the Celtic Shorthorn skulls agree with the skull of an 



SKULLS OF OXEN' FROM NEWSTEAD. 



263 



Ox from Irkutsk figured by Duerst *. In cross-bred cattle the 
preniaxilla may be forked on the one side, as in text-fig. 73, but 
long and rounded on the other as m text-fig. 71. 

Text-fi?. 75. 4 




Front part of skull of a honied Cadzow Ox with notched premaxilla; (P.M.) which 
fail to reach the nasals (Na.). From a skull in the Roj'al Scottish Museum. 

Text-fig. 76. 




Front part of skull of cross-hred Shorthorn in which the premaxilhe (P.M.) 
fail to reach the nasals (JYct.). 

As alread}^ mentioned, Dr. Gray stated in the British Museum 

* Wilckens, ' Naturgeschichte der Haustiere/p. 305,1905. 

18* 



264 PROF. J. C. EWART ON 

Catalogue of the Ungulata that in the true Oxen " the premaxillse 
are large and always extend upwards between the maxillae and 
nasals and consequently articulate with both these bones as in 
Buhalus.'" Lydekker, in discussing the premaxillfe in the Bovidae, 
points out that in Bos chinensis, a new species desciibed by Gray 
in 1870 (L e. before the pxxblication of the Catalogue of the 
Ungulates), "the premaxillse are small and do not extend upwards 
to the nasals precisely as in Bibos"*. But while in Bos nania- 
dicihs and in Bos chinensis the premaxillae may be always small, they 
are not invariably small in the Bibovine group. In a Gaur skull 
in the Royal Scottish Museum the premaxillse reach the nasals, 
and Mr. Pocock informs me that, of four Gaur skulls he examined 

Text-%. 77. 




Skull of polled Ne^vstead Ox with forked premaxilte, large orbits, deeply notched 
occiput, uneven forehead ending in a rounded mesial prominence. Skulls of 
this type with horns are sometimes said to belong to JBos frontosus Nilsson. 

in the British Museum, one has the premaxillae in contact with 
the nasals, while in another skull they are separated by a long 
interval from the nasals. Of all Lydekker's statements about the 
premaxillse, the one which bears most on the present enquiry is to 
the effect that a small premaxilla is " never found in any European 
Ox " t. If the premaxilla is small in Bos oiamadicus, but never 
small in European cattle, the presumption is that European cattle 
are not descended from Bos namadicus, biit from Bos j^riinigenms, 
in which the premaxillse in all the skulls examined reach the 
nasals. 

In one of theNewstead skulls, which probably belongs to an im- 
ported Ox, as large as a modern Aberdeen- Angus steer, the total 
length of the premaxilla is only 118 mm. — 47mm. shorter than the 

* Memoirs of the Geological Survey of India, series x. vol. i. p. 19. 
f Oj). cit. p. 3. 



SKULLS OF OXEX FROM NEWSTEAD. 



265 



premaxilla of the Aberdeen-Angus heifer represented in text-fig. 
71. The distance between the premaxilla and the lachrymal in 
this Newstead Ox is 80 mm., in the Aberdeen- Angus only 25 mm. 



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But in addition to being small, or rather because it is .small, it fails 
to reach the nasal and makes no attempt to fill up the gap between 



266 



PROF. J. C. EWART ON 



the maxilla and the nasal (text-fig. 74). As it happens, there is 
a skull of one of the so-called " wild " white Cadzow cattle from 
Hamilton Park in the Royal Scottish Museum Avhich probably 
belonged to an Ox including the Celtic Shorthorn amongst its an- 
cestors. In this Ox (text-fig, 75), though the premaxilla is 142mm, 
in length it neither reaches the nasal nor yet extends into the gap 

Text-fig, 79, 




F.M. 



F.M. 

Outlines of tlie occiput and part of the forehead of three Urus skulls. 



J?}\, frontal ; F.3I., upper border o( foramen magnum. 
In all three skulls the iutercornual ridge projects far beyond occipital condyles. 
From skulls in the Anatomical, Antiquarian, and Free Church College 
Museums, Edinbiirgh. 



between the maxilla and nasal. In shape and in being notched at 
its proximal end, it reminds one of the premaxilla of the Celtic 
Shorthorn (text- fig. 73). Another skull with premaxillte which fail 
to reach the nasals may be mentioned, because it has many of the 



SKULLS OF OXEN FROM NEWSTEAU. 



267 



characteristics of the Urus. In tliis skull (text-fig. 76), which 
belonged to a. cross-bred Shorthoi'ii, the })reiuaxillix in shape closely 
agrees with the preniaxilki of the Urns, but it is i-elatively shorter 
and neither reaches the nasal nor extends into the space between 
the nasal and the maxilla. 

Text-fio-. 80. 




Front view of skull of Bison (Bos bison). 

In the Bison the horn-cores arc near the orbits, the parietals (Pa.) are not o^•erlappcll 
by the frontals (Fr.), and tlic preniaxilla3 (P.M.) are far removed from the 
nasals (Ka.). 

Though very few skulls have been available for study, it is 
evident (1) that in some modern Europeancattleas wellasin some 
of the cattle in Britain during the Roman occupation, the pre- 
maxillai were short and failed to reach the nasals, and (2) that as 



2G8 PROF. J. 0. EWART OX 

the premaxillfe vary in wild as well as in domestic cattle the relation 
of the premaxillfe to the nnsals is of little diagnostic value. 

The Forehead.— In the NeAvstead skulls the forehead varies 
considerably ; in some specimens it is flattened as in a typical Urus 
(text-fig. 66), and the ridge between the horn-cores is nearly 
straight ; in others there are prominences and depressions and the 
intercornual ridge is arcuated as in some specimens of the Ui'us, 
Avhile in a polled'skull (text-fig. 77) of the/ro«ios^r.s type, the vertex 
projects forwards and npwai'ds to foi-m a well-marked rounded 
mesial prominence. In this polled skull there is also a projection 
from the middle of the forehead and a well-marked ridge at each 

Text-fig. 81. 




Hind part of skull of tlie Urus represented in text-fig. 66. 

The front of tlie horn-core is in a line with the occipital condyle, and the temporal 
fossa IS closed behind by a plate of bone which supports the horn-core. 

side between the orbit and the long deep orbital sulcus. In old 
animals the parietals seem to be completely covered by the 
frontals. But even when the intercornual ridge pi'ojects fax- 
l)eyond the level of the occij^ital condyles the interparietal 
proljabl}" enters into the formation of the forehead. That the 
parietals (or the interparietals) form part of the forehead in polled 
Oxen is indicated b}^ text-fig. 78 B, J^a. That in horned Oxen the 
parietals al.so enter into the formation of the forehead is suggested 
by a large skull of the Urus tyjse in the Royal College of Surgeons- 
Museum, London (No. 1121 A). In this skull the frontals are 
separated by a wedge-sliaped piece of bone (probably the inter- 



SKULLS OF OXEN FROM XBWSTEAD. 



2()f> 



parietal), which forms the middle portion of the intercornunl 
ridge. Oscar Schmidt * states that in a. front view the ptirietals 
" cannot be seen at all," bnt probably in all cases the parietals 
in the Ox, as in the Bison (text-fig. 80), enter into the formation 
of the forehead. The great specialization of the frontal region of 
the skull in the true Oxen is made evident by the examination of 
f(otal skulls. In a very young foetus tlie frontals (text-fig. (58, Fr.) 
only form the fi'ont portion of the I'oof of the cranium ; but about 

Text-fio. 82. 





rrontlct and lioni-coros of a small Newstead Ox with a prominent forehead. 
The horn-coreK curve upwards and backwards. 



the fifth month they ai'e relatively very much larger than the 
parietals (text-fig. 69, Fr.), and at birth they form nearly the entire 
roof of the cranium (text-fig. 70). After birth the frontals 
continue to grow back\\ards, and they eventually form all but the 
wedge-shaped middle portion of the intercornual ridge. In the 
Urus the frontals sometimes project 100 mm. (4 indies) beyond 
the occipital condjdes (text-fig. 79), but in the Aberdeen-Angus 
the occiput almost forms a right angle with the frontals, and the 

* 'The Mammalia,' p. 170. 



270 



PKOF. J. C. EWART ON 



parietals (text-fig. 78 B), as in the Bison (text-fig. 80), obviously 
enter into the formation of the forehead. The highly specialized 
condition of the occipvit in the Uvus has resulted from the horns 
having migrated upwards from their original position above the 




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orbits (text-fig 63) to lie entirely behind a line passing through the 
occipital condyles (text-fig. 81). In addition to Newstead skulls 
with the foi'ehead flat or presenting lidges or prominences, thei'e 
is a skull (text-fig. 82) in which the forehead is so convex that the 



SKULLS Ui-' UXEX jb'llUM XEW.SXEAD. 



271 



horn-cores at their origin are far behind tlie most prominent part 
of the forehead. jVEoreover, the horn-cores in this case are flat- 
tened and curve outwards and Ijackwai-ds as in some Eastern 
cattle. In their position and direction the horns in this Newstead 
Ox decidedly differ from the horns (also of the flat type) of a 
Shetland heifer in my possession, which project almost directly 
upwards as in some Zebus. 




The horn- cores are, as a rule, circular in section and, in skulls of 
the Celtic Shorthorn type, curve outwards and forwards nearly 
at a right angle to the frontals. In other skulls they curve out- 
wards and upwards, or outAvards and downwards, or outwards, 



272 



PROF. J. C. EWART ON 



upwards, and backwards as in some Indian cattle. Excei:)t when 
they curve forwards at right angles to the frontals, as in typical 
Celtic Shorthorns, the horns assist but little in settling the I'ace 
to which the Newstead skulls belong. 

The Occijmt. — In some ISTewstead skulls the occiput has the 
characteristics of the Urus represented in text -fig. 83. The general 
outline is quadrangular, the occipital crest is flattened but not 
encroached on by the tempoi'al fossae, the latei'al boitlers are nearly 
straight, and a line carried through the highest point of the crest 
lies beloAv the centre of the horn-cores ; further, there is a shield- 
like projection [L.X.) for the insertion of the ligamentum n\u-ha^. 
The distance from the crest to the lower border of the foramen 
niagnum is nearly the same as the distance between the condyles 
(C.) and little more than half the distance between the notches 
(^V.i\^.) below the horn-cores on a level with the temporal fossfe. 

. Text-fis-. 85. 




Occiput and lioni-coies of a Urus in which the notches below the horn-cores 
are deeper than in text-fig. 83. 

This occiput closelj' resembles the occiput of a Bos namadicus skull 
in the British Museum. 



The nearly smooth supracristal part of the occiput extending 
between the horn-cores measures from above downwards about 
half as much as the part lying between the crest and the lower 
border of the yb?r«»e9i OT,o(/?ww?z. In some cases the supracristal 
part is flat and nearly in a line with the occiput proper, in others 
it projects beyond the crest (text-fig. 79) and, though concave 
in the centre immediately over the shield for the ligamentum 
nuchse, it is prominent and convex above and forms a marked 
angle with the forehead. In some cases the upper border of the 



SKULLS OF OXEN FROM NEWSTEAD, 273 

occiput viewed from behind is nearly straight, in otliers it is 
slightly concave or distinctly arcuated. 

In specimens of the TJvus with veiy thick horn-cores a bi-idge of 
bone extends downwai-ds from the base of the horn-core a,nd forms 
a posterior wall for the temporal fossa (text-figs. 81 &■ 83, y.), which 
completely cuts off' the fossa, from the occiput. It is owing to the 
temporal fossa? being closed in behind by extra supports for the 
horn-cores that the width of the occiput immediately below 
the horn-cores is sometimes nearly twice as much as the distance 
between the crest and the lower border of the foramen magnum. 

Though in the majority of the !N"ewstead skulls the occiput 
conforms to the Urus type there are several with short premaxilla? 
in which the occiput reminds one of Lydekker's Bos acutifrons. 
Writing in 1880 Lydekker pointed out that Bos acutifrons of the 
Punjab Siwaliks had no marked I'elationship to any existing species 
and was " widely difiei-ent in the form of its occiput, frontals, and 
horn-coi'es from Bos 2)rimigen,ius" * . 

Though Riitimeyer and others vegA.Yy{Q({ Bos 2)lanifrons and^os 
acutifrons iia varieties of a species closely allied to Bos primigenms, 
no skulls of true Oxen have hitherto been described in which the 
occiput resembles that of Bos acutifrons. 

In Bos Qiamadicus, SiS well as in Bos 2^)' imi genius, the notches 
below the bases of the horn-cores are shallow, but in the jS!"ewstead 
skull represented in text-fig. 84 indentations or notches (iV".) 
below the horn-cores are so deep that the connection between the 
upper (parieto-frontal) part of the occiput and the lower (infra- 
cristal) part is relatively short. Because of these indentations the 
occiput viewed from behind (text-fig. 84) bears a resemblance to 
that of the Gaur (Bos gaurus). 

In Bos primigenius the summit of the occipital cxest is on a 
lower level than the middle of the horn-cores (text-fig. 83), 
but in the Newstead skull of the acutifrons type a line carried 
through the summit of the rounded arch formed by the occipital 
crest (text-fig. 84) passes above the middle of the horn-cores. 

At the widest part the occiput of the first century skull 
(text-fig. 84) measures 215 mm., and from the lower border of 
the foramen magnum to the occipital crest the distance is 
115 mm. The depth (115) multiplied by 100 and divided by 
the width (215) gives an index of 53"5. In the Urus represented 
in text-fig. 85 the corresponding index is 47. Between the 
notches under the horn-cores the distance in the Newstead 
skull (text-fig. 84) is 123 mm.: 123x100-^-215 (the total 
width of occiput) gives an index of 57 ; in the Urus, owing to the 
shallowness of the notches under the horn -cores, the correspond- 
ing index may be 90. The distance (115 mm.) between the 
lower border of the foramen magnum and the occipital crest 
multiplied by 100 and divided by the distance (123 mm.) between 
the infi-acornual notches gives an index of 93*5 ; the correspond- 
ing index in the Urus may be only 52*4. 

* Memoirs of tlie Geological Survej^ of India, series x. vol. i. p. 2. 



274 



PKOF. J. C. KAVAllT ON 

Text-fie-. 86. 




A. — Front view of the Newstead skull of which the occiijut is represented in 

text-fig. 84, p. 271. 

B. — Temporal fossa of above skull. The fossa communicates freelj' with the 

occiput. Compare with fossa of the Urns (text-fig. 81, p. 268). 



SKULLS UF OXEX FROM XKWSTKAD. 

Text-lii--. 87. 





A.— Front view of a wliito " wild " Cailzow Ox from Hamilton Park. 
B. — Occiput and liorn-corcs of the above skull. 
This skull closely resembles the Newstead skull (text-figs. SJ; & 86) with a (1eepl\ 
notched occiput. From a specimen in the Royal Scottish Museum. 



276 



PROF. J. C. BWART OX 



From the naiTOw shield-shaped projection {L.X.) vmder the 
crest (text-fig. 84), a narrow ridge extending towards the foramen 
magnum divides the lower part of the occiput into two equal 
portions x-oughened for the attachment of muscles. Between the 
crest and the upper border of the foramen magnum the occiput 
is distinctly concave — near the crest it is excavated to a depth 
of 12 mm. The part of the occiput external to the condyles {C.) 
is divided by a ridge continuous with the incurved paroccipital 
processes {P-P-) into two irregidar, more or less concave, rough 
surfaces. The greatest width across the occipital condyles {€.), 
is 99 mm. (only 23 mm. less than the space between the 
infracornual notches): 99x100-h123 (the width of the occiput 
under the horn-cores) gives an index of 80. The corresponding 
index in the Urus skull (text-fig. 83) is only 55. 

. Text-fio-. 88. 




Occiput and liorn -coves of an American Bison. 

The frontals neitliev cover the parietals (text-tig. 80, p. 267) nor project beyond the 
level of the occipital crest, and the horn-cores arc separated by wide gaps 
from the supra-occipital. 

The part of the occiput above the occipital crest is divided into 
three portions, \\z. : — a mesial semicircular deeply excavated 
portion and two lateral portions continuous with the horn-cores 
externally and the forehead above. The mesial semicircular 
concave portion (text-fig. 84), 60 mm. wide and 20 mm. deep, has 
a rough margin, 3 to 6 mm. in thickness, which forms the nearly 
horizontal border of the mesial frontal protuberance (text- 
Hg. 86, A). 

From the measurements given it follows that the occiput of 



SKULLS OF OXEX FROM XEWSTEAD. 277 

the Newsteacl skull (text-fig. 84) differs pi-ofounclly from that of 
the more highly specialized Urus skull represented in text-fig. 83. 

The Horn-cores. — The horn-cores of the Newstead skull, 
characterized by a deeply notched occiput, are pyriform as in 
Bos aciitifrous — they measure at their origin 60 mm. from above 
downwards and 45 mm. from before backwards. At a distance 
of 30 mm. fi-om the beginning of the gi-ooved surface they 
measure 56 mm. by 42 mm. The fragments of the horn-cores 
present (text-fig. 86, A) are convex above, grooved in front and 
below, and extend outwards and slightly forwards. 

The Teriijwrcd Fossa.^ln the Urus skull figured the temporal 
fossa is completely closed behind (text-fig. 81) by a wall of bone 
which helps to support the horn-core (text-fig. 83) ; the external 
opening of the fossa, rounded and contracted behind, is pointed 
in front and arched above (text-fig. 81). In the Newstead skull 
with short premaxillse and a notched occiput, the temporal fossa 
is wide behind, where it opens on to the occiput, wider than in 
the Urus in front, and the upper border is sinuous (text-fig. 86, B) 
as in the Bison. 

The Base of the Cranium. — In the Urus the anterior as well as 
the posterior tubercles are well developed, but in the Newstead 
skull (text-fig. 84), as in Bos acutifrons, the anterior tubercles are 
small and inconspicuous. 

In having the lower jjart of the occiput separated from the 
iipper by deep notches ujider the horn-cores the Newstead skull 
(text-fig. 84) decidedly diflers from Bos primigenias (text-fig. 83) 
and also, though to a less extent, from Bos namadicus. Again, in 
having the occipital crest overhanging the true occiput, the 
Newstead skull differs from the Gaur and Banting, 

If Lydekker is right in assuming that the occipital crest in 
Bos acutifrons " extends upwards to within a short distance of 
the vertex cranii so that the supra-cristal portion of the occipital 
region is reduced to a very narrow band '' *, the Newstead skull 
also differs from Bos acutifrons. If, however, in Bos acutifrons 
the mesial depression immediately below the vertex is supra- 
cristal, the Newstead skull with a deeplj' notched occiput may be 
regarded as belonging to a race allied to, or descended from. Bos 
acutifrons of the Punjab Siwaliks. 

It will doubtless be asked. Are any of the modern breeds of 
cattle characterized by short premaxilla? and an occiput of the 
primitive type represented in text-fig. 84 ? As it happens, the 
skull (text-fig. 87) of the white " wild " Cadzow Ox with short 
premaxilhe (text-fig. 75) has an occiput (text-fig. 87, B) of the 
acutifrons type, and otherwise resembles several of the Newstead 
skulls. The skull of the Cadzow Ox seen from behind looks 
almost as if it were intermediate between the Bison (text-fig. 88) 
and a Urus with fairly deep notches (text-fig. 85). 

At one time the Hamilton Park herd of white cattle consisted 

* Memoirs of the Greological Survey of India, ser, x. vol. i. p. *J7. 

Proc. Zool. Soc— 1911, No. XIX, 19 



278 



PIUJF. J. C. EAVART ON 



almost entirely of polled individvials, now they are said to be all 
horned *. There is a tradition that Oadzow cattle re-acquired 
horns through a Highland bull, which, for a time, took forcible 
possession of the herd, and it is a matter of history that some 
years ago a " wild " Chillingham bull was made use of to rein- 
vigorate the Cadzow " wild " cattle. The intercrossing, perhaps, 
led to reversion towards a I'ace in the possession of, if not 
actually introduced by, the Roman auxiliaries who garrisoned the 
border-fort during the later part of the first century t. 

Text-fig, 89. 




Front part of the skull of a Zebu {Bos indicus) in which the pi-emaxillaj 
reach the nasals. 



In addition to Newstead skulls with an occiput of the Urus 
and acutifrons types, there are skulls which in the occiput 
agree with certain Indian cattle. In Indian as in European 
domestic cattle the premaxillse are sometimes long and in 
contact with the nasals (text-fig. 89), sometimes shoit and 
terminating some distance from the nasals (text-fig. 90). In 
at least some Zebus long premaxillsB ai'e correlated Avith a wide 
Urus-like occiput, and short jDiemaxillse Avith a narioAv deep 
occiput. In seA"eral small NeAA^stead skulls the occiput closely 

* A skull of a polled Cadzow Ox in the Anatomical Museum of the University 
of Edinburgh agrees in the premaxillse and occiput with JBos frimigenms. 

t The Chillingham and Chartley " wild " park cattle in their occiput conform to 
the Urus type. 



SKULLS OV OXEN FROM NBWSTEAD. 



279 



agrees with a Zebu skull (text-fig. 91) in the Royal College of 
Surgeons Museum, London. In Bos 2)rimigeniics the shield-like 
projection for the ligameutum nuchce is in contact with the 
occipital crest, but in some of the small Newstead skulls, as in 
the skull of the acutlfrons type, the rough surface for the attach- 
ment of the ligameiitimi nachce lies, as in some Zebus, nearly 
midway between the occipital crest and the upjjer border of the 
foramen magnum. Further inquiry may show that Indian 
domestic cattle are in part descended from ancestors allied to 
Lydekker's Bos acatifrons and that the Celtic Shorthorn [Bos 
lojtgifi'ons Owen, Bos hrachyceros Riitimeyer) is intimately related 
to some of the small Oriental races. 

Text-fig. 90. 




Front part of the skull of u Zebu in wliicli the premaxilltie fail to reaeli 
the nasals. 



The Origin of the Newstead Oxen, 

In addition to cross-bred iinimals Newstead has yielded five 
fairly distinct types of Oxen, viz. : — (1) Oxen of the Celtic Short- 
horn type ; (2) Long-horned Oxen of the Urus type ; (3) Oxen 
with an occiput of the Bos acutifrons type ; (4) Oxen with a 
convex forehead, an arcuated intei'cornual ridge and horns curving 
outwards and backwards : and (5) Hornless Oxen. It is im- 
possible to say definitely how any of these forms originated, but 
it may be safely assunred that they were not all foi-med in 
Europe from Bos ■priniigenias. 

19* 



280 



PROF. J. C. EWART ON 



Though Bos j)rimigenms reached Europe from Central Asia in 
early Pleistocene times, there is no evidence that it was living 
under domestication on the arrival in Post-Pleistocene times of 
the Neoliths. Further, there is no evidence of the existence of a 
small Ox of the Celtic Shorthorn type in Eui'opean Palaeolithic 
deposits — Prof. Boule e. g., found not a single fragment of Owen's 
Bos longifrons in deposits of the Reindeer age at Monaco. 
Neither is there any evidence of the existence of a small wild Ox 
in Pleistocene times in Central Asia. 

Text-fig. 91. 




Occiput of the Zebu with the premaxilke reaching the nasals (text-fig. 89). 

In several of the Celtic Shorthorn skulls from Newstead the occiput resembles 
that of the Zebu. 

Text-figs. 89-91 from skulls in the Royal College of Surgeons Museum, London. 



The examination of the bones of Oxen from Anau, Tui'kestan, 
led Duerst to conclude : — (1) that a large long-horned breed was 
formed by the Anau.-li about 8000 B.C. from a large wild Asiatic 
race which he regarded as the exact equivalent of the European 
TJrus (Bos prlmigenius) ; (2) that about 6000 B.C. a small short- 
horned breed, identical with Owen's Bos longifrons and Riiti- 
meyer's Bos bracJiyceros, was formed at Anau, or brought to Anau 
from some other settlement in Centival Asia. If, as seems 
probable, the Urus was the only wild Ox in Central Asia in 
prehistoric times, it must be assumed that the small Ox in the 



SKULLS OP 0\BX FROM NEWSTEAD. 281 

possession of tlie Anau-li was a dwai-fed descenrlant of an Asiatic 
variety of Bos jjrimigenius *. 

It is doubtless possible that a small breed may have been 
formed out of the huge Urus by the ISTeoliths immediately after 
they reached Europe, but the evidence, so far as it goes, suggests 
that the Neoliths brought the "Celtic" Shorthorn with them 
from Central Asia. 

Though in Britain the Urus was hunted by the Neoliths — 
evidence of this we have in the Urus skull from Burwell Fen, 
near Cambridge, with the frontals pierced by a Neolithic flint 
implement — thei-e is no evidence that Bos primigenius was once 
domesticated in Britain or that the Neoliths allowed their 
domestic cattle to breed with young wild bulls. 

But on the Continent the Urus was apparently domesticated at 
a comparatively early period and crossed with the small breed 
originally brought from Central Asia. Hence it may be said 
that up to at least the Bronze age the majority of the domestic 
cattle in Europe were the descendants of Bos primigenius — some 
being nearly pure descendants of the imported " Celtic " Short- 
horn breed, while others were pure or nearly pure descendants 
of the indigenous wild Urus (Bos taurtis jjrimige^iitis). 

There is no evidence that there existed in Europe or in Central 
Asia a variety of Bos j)rimigeniiis with the occiput deeply notched 
and otherwise resembling the one represented in text-fig. 84. 
Neither is there any evidence that in Bos namadicus — the Urus 
of India — the occiput was deeply notched or characterized by an 
excavated intercornual lidge. The only extinct form to which 
the Newstead skull represented in text-figs. 84 and 86 bears any 
marked resemblance is Bos acuiifrons of the Pliocene Siwaliks. 
It has been suggested that the Newstead skull with a deep mesial 
semicircular depression above the occipital crest belonged to a 
hybrid between an Ox and a Bison, but this view is not supported 
by the skulls of Ox-Bison hybiids. Moreover, in its occiput and 
premaxillfe this Newstead skull (text- figs. 84 & 86) very closely 
agi-ees with the skull of a Cadzow Ox (text-fig. 87) in the Royal 
Scottish Museum. It may hence in the meantime be assumed 
that some of the cattle in the south of Scotland during the 
Roman occupation were descended from an Indian race allied to 
Bos acutifrons. 

Of the Newstead cattle with horns curving backwards and down- 
wards (text-fig. 82) it need only be said that they seem to be more 
intimately related to Bos namadicus than to Bos prmiigeniiis. 

The polled Newstead cattle represent two distinct types. Some 
had a neai^ly flat forehead, a nearly straight " intei-cornual " ridge, 
and a square-shaped occiput; in others the forehead was very 
uneven and ended in a pronounced mesial prominence which 
projected upwards and forwards (text-fig. 77). The Newstead 

* The small Ox of Aiiau is probably now represented in Asia bj' the long-browed 
Zebus characterized by small horns of the Bos longifrons type. 



282 MR. G. P, FARBAN ON COPEPODA 

cattle with a flat poll (as in the modern Galloway and in polled 
Cadzow cattle) obviously belong to the Urus type, while those 
with a mesial prominence seem to belong to the frontosus type of 
ISTilsson. As there were hornless cattle in Egypt as early as the 
Fourth Dynasty, we are not likely soon to ascertain where or 
Avhen polled breeds originated. There is no evidence that any 
of the varieties of polled cattle are descended from hornless wild 
ancestors; neither is there certain evidence of a pure horned 
race suddenly producing polled offspring. If, as seems highly 
probable, the absence of horns is not due to reversion, the polled 
condition was either acquired by the gradual reduction in the size 
of the horns or suddenly, ?'. e. by mutation. In polled breeds 
which now and then produce individuals with " loose horns " the 
polled condition Avas perhaps acquired slowly, while in polled 
breeds in which " scurrs " are unknown the polled condition was 
perhaps due to a mutation. Though the Galloway and the white 
polled " wild " Cadzow cattle may have descended from Bos 
primigenius, the white polled Somerford, and the round polled 
Aberdeen- Angus cattle with the premaxillfe extending well vip 
between the maxillse and nasals, may be the descendants of an 
Oriental race allied to a modern Syrian breed apparently in the 
act of losing the horns. 



13. Plankton from Christmas Island, Indian Ocean. — I. On 
Copej)oda of the Family Corycmdce. By George P. 
Fare AN *, 

[Received Octoljev 31, 1910 : Read February 7, 1911.] 
(Plates X.-XIY. t) 

Through the kindness of Dr. W.T. Caiman I have had theoppor- 
tunity of examining the Copepoda of a small collection (8 bottles) 
of Plankton, made by Sir John Murray, K.C.B., F.R.S., and 
Dr. C. W. Andrews, F.R.S., at Christmas Island in the Indian 
Ocean, and presented to the British Museum by Sir J. Murray. 

The gatherings were all made at approximately the same time 
(July- August, 1908) and in the same locality, on the north side 
of the island in shallow water neai- shore, and for this reason 
it has not been thought necessaiy to refer to each of them 
separately. 

The collection, though small in bvilk, is exceedingly rich in 
species, and the genus Corycceus is especially well represented. 

It has been recognized that there are, in the the genus Cory- 
cceus, two groups differing from each othei' in several distinct 
characters, the most notable being the form of the ventral 
process, situated between the maxillipedes and the fii-st pair of 

* Communicated by Dr. W. T. Calman, F.Z.S. 
t For explanation of the Plates see ^. 296. 



P Z.S.19I1. PI. X. 




Hatk,l.itli'. Loudo 



1 6. CORYCELLA B RE VIS 

7 11. COKYCELLiV CURT/v. 



p. Z.S.1911.P1XI. 




1-6. CORYCPJ.T.A. CTIRT.^. 7. C.BKEVIS 
8,9, GOHYCE.LLA sp. c? ]0. G.CARTNATA. 
11,12. CORTCAEUS GRi^CILTCAUDATUS 



PZ.S.1911, Pl.XTT. 




1—3. CORY"CAEUS CAT US ^_ 6. C OBTUSUS 

7. C. DTJBIUS. 8._9, C. TENUIS, 



p. z.s. 1911. PL xrn. 




1_6. CORYCAEUS MURI^AYI. 
7_9. C. ANDREWSI. 



Hutli^Iiitii^ L oudon. 



p. Z.S.1911.P1.XIV. 




1_^. CORYCAEUS AKTDREWSI. 
5_a. C. DUBIUS. 



Hatli Jattf London:. 



FROM CHRISTMAS ISLAND. 283 

feet of the female, which is beak-shaped in one group and semi- 
circulax' in the other. There can be no doubt tliat these characters 
are, collectively, of generic importance, and I propose the name 
Corycella for that subdivision, as typified by C. gihbulus, which 
is marked by the possession of a beak-shaped ventral process, 
leaving the name Corycoeus for the remaining sj)ecies, of which 
O. speciosus may be selected as the type. 

The genus Corycams was founded by Dana in 184.5 (6), his 
diagnosis being repeated in 1846 (7), but no species belonging to 
the genus was described till 1849 (8). Dana's definition of the 
genus includes both the genera defined above, and his papers 
describing the species, first published without figures in 1849 (8), 
and afterwards with figures in 1852 (9), contain forms belonging 
to both Corycella and Corycceus, but do not indicate any one of 
them as forming the type of the genus. 

The principal differences between Cwycceus and Corycella may 
be summarized as follows : — 

Genus Corycella, nov. Genvis Coeyc^us Dana. 

Abdomen of <? and $ one-jointed ; Abdomen of (J and $ usually two- 

4th thoracic segment without lateral jointed ; 3rd and 4th thoracic segments 

points, not distinct from 3rd thoi'acic distinctly separate, the latter with 

segment. Ventral process beak-shaped lateral points. Ventral process semi- 

in ^ . 2nd antenna with setae on 1st and circular in ^ . 2nd antenna with the 

2nd basal joints distantly feathered ; setae on 1st and 2nd basal joints not 

terminal spine short in ^; inner edge of feathered; terminal spine elongated in 

2nd basal coarsely feathered. ,J> inner edge of 2nd basal usuallj' with 

one or more strong teeth. 

1st to 3rd feet, exopodite with 0.0.1 1st to 3rd feet, exopodite with 1.1.3 

outer edge seta;. outer edge setae. 

4th foot without endopodite, exopo- 4th foot with endopodite bearing one 

dite with 0.0.1 outer edge setae. or two setae, exopodite with 0.1.1 outer 

edge setae. 

The one- jointed abdomen and the feathered setse of the second 
antennae occur, as Dahl has pointed out, as characters of im- 
matu.rity in the genus Corycceus. 

Giesbrecht (10) recognises five species of Cory cams in the 
group for which the name Corycella is proposed, viz., C. gihhulus 
Giesbr., C. concinnus Dana, C. rostratths Glaus, C. carmatus 
Giesbr., and C. longicaudis Dana.' In Dahl's revision of the 
genus (5) he accepts C. gibbuhis, C. concinnus, C. carmatus, 
C. rostratus, and, under the new name of C. tenuicattda, C. longi- 
caudis, asserting that Dana's description of C. longicaudis really 
refers to C. S2ieciosus juv., a view which is strongly supported 
by Dana's figures. Dahl also recognises Dana's (8) description of 
C. gracilis as being good, but does not give any reason for this 
opinion. Dana's description and figures of G. gracilis evidently 
refer to a male of Corycella, but the specific differences between 
the males of this genus are so slight that it does not appear 
possible to refer the description and figures with certainty to any 
particular species. Dahl's belief that G. j^elhicid^is of Dana is the 
female of what he recognises as C. gracilis also appears to be 
a mere conjecture, there being very little evidence for or against 



284 MB. G. p. FARBAN ON COPEPODA 

tlie view in Dana's figures, which, apparently, include two 
different species. 

Wolfenden (11) also believes that he has recognised the 
C. 2}6ttit'Cidus of Dana, but his figures and descriptions refer 
undoubtedly to the species which Giesbrecht has described as 
C. gibhidus. 

Dahl, in the paper referred to above (5), which is a prelimi- 
nary note on the Oopepoda of the Plankton Expedition, has 
drawn up a useful diagnostic table of the members of the genus 
C'oryccBus as recognised by. him. The poi-tion which refers to the 
genus Gorycella has been translated by Dr. Wolfenden (11) in his 
account of the Maldive Oopepoda. In this table Dahl gives as 
a mark to distinguish the other members from C. rostrattis, 
"Yiertes Beinpaar mit einem inneren Za pf en welch er eine Borste 
triigt." If by " Zapf en " is meant endopodite, as is undoubtedly 
the case in the latter part of the table, and as Wolfenden trans- 
lates it, this statement is an error, as the endopodite is absent 
throughout the group. There is a misti-anslation in Wolfenden's 
table which detracts somewhat from its usefulness ; the sentence 
" Genital openings of $ set at end of abdomen " should read 
" distant from end of abdomen," " vom Ende des Abdomens 
entfernt." 

The specific characters in the genus Gorycella are very slight 
and depend mainly on the shape of the abdomen and the third 
thoracic segment. The following is an attempt to draw up 
a diagnostic table, to include the females of which a recognisable 
description exists and two new species from the Christmas 
Island collection, which are describee! below. 

1. Furca only twice as long as broad C. rostrata. 

2. Fnvca three or more times as long as broad. 

A . Furca shorter than widest part of abdomen in dorsal view. 
a. Widest part of abdomen behind the middle, 3rd thoracic 

segment with dorsal hvimp. 
a. Abdomen less than twice as long as high in lateral 

view, median furcal spine short and stout C. f/ilihila. 

/3. Abdomen ca. 2f times as long as high in lateral view, 

median furcal spine slender C. Irevig, sp. n. 

h. Widest part of abdomen in front of middle, 3rd thoracic 
segment not humped dorsallj'. 
a. Thoracic spines long, abdomen with ventral setose 

pad C. ca rinata . 

13. Thoracic spines short, abdomen without setose pad ... C. citrta, sp. n. 

B. Furca longer than widest part of abdomen. 

a. Furca less than half as long as abdomen, 3rd thoracic 

segment with dorsal hump C. Inngicmidis. 

b. Furca less than half as long as abdomen, 3rd thoracic 

segment without dorsal hump C. concinna. 

CoRYCELLA GiBBULA Giesbr. 

Coryccmis gihhidus Giesbrecht (10). 

Corycmus pellucidus Wolfenden (11). 

? Corycceus onegalops Will.-Suhm (1). 
Common in all the gatherings, upwards of 120 specimens. 
The most easily recognised characters of the female of this 



FROM CHRISTMAS ISLAND, 285 

species are tlie deep ceplialon with eyes rather small and set far 
apart, the abdomen widest at its posterior third (without furca) 
and usually bearing two spermatophores, the furca contained 
2t times in the rest of the abdomen, the middle furcal seta thick 
and equal in length to the width of the furca, and the setose pad 
on the antero-ventral pai-t of the abdomen. 

Wolfenden (11. p. 1027) has described at some length a female 
Coryca'us which occurred plentifully in the Maldive collections, 
and which is apparently identical with the above. He ascribes 
it to G. peUucidus Dana, and records C. gibbulus as a separate 
species. He remarks, however, that he believes that the C. pel- 
htcidus of the IMaldive collections is probably identical with 
0. gibbulus. His figure of the fourth foot (fig. 9, pi. xcix.) is 
evidently incorrect in showing an endopodite bearing a long seta, 
as in the text it is said that no endopodite is present. 

Giesbrecht's (10) figure of the female abdomen of C. gibbulus in 
lateral view is not so deep nor so irregular in outline as are all 
my specimens ; in this they agree more nearly Avith Wolfenden's 
figure, but I think thei-e can be no doubt that all these forms are 
identical. 

It seeins ]iot improbable that Willemoes-iSuhm's C. onegalops 
was in reality this species, but his description and figure, as 
given by Brady (1), can hardly be regarded as constituting a valid 
diagnosis. Bi'ady's C. peUncidtcs may also include it with others. 

Distribution. Red Sea, Arabian Sea, Indian Ocean (many 
recoiTls), and Tropical Pacific. 

CORYCELLA BREVIS, Sp. n. (PI. X. figS. 1-6, PL XI. fig. 7.) 

Female (PI. X. figs. 1, 2). — Length -85 mm. ; cephalothorax, 
in middle line, '54 mm. ; abdomen and furca -3 mm. ; cephalon 
rather deep, curved doi'sally in lateral view ; eyes large, pro- 
minent in dorsal view. 

2nd thoracic segment with slight dorsal hump, as in C. gihbula 
and ( \ longicmtdis. 

3rd thoracic segment (PL X. fig. 6) with comparatively short 
lateral points. 

Abdomen widest at its posterior | excluding furca ; in lateral 
view the abdomen is parallel-sided for about 4' of its length, and 
is then tapered to the furca ; it has a slightly spinulose antero- 
venti'al boss. 

Fui"ca about 6 x 1, contained 21 times in rest of abdomen ; its 
median terminal spine not very strong, about twice as long as 
width of furca. 

1st and 2nd antennse (PL XI. fig. 7) as in C. gibbula. 

1st to 3rd feet (PL X. figs. 3, 4) .almost the same as in C. gihbula. 
The 3rd joints of the exopodites of the 1st .and 2nd feet are 
finely serrate, in the 3i-d foot the serrations are not visible. 

4th foot (PL X. fig. 5) as in (■. gihbida. hollowed on distal 
margin of 2nd basal ; inner margin of the 2nd basal forms an 
acute angle with its base. 



286 MR. G. p. PARRAN ON COPEPODA 

The characters which distinguish this species from C. gibbula 
are the proportionately greater length and different form of the 
abdomen and the shorter 3rd thoracic spines. 

Only one specimen was found. 

CoRYCELLA coNCiNNA Dana. 

In five gatheinngs, about 30 specimens. 

Giesbrecht (10) has given a figure of the female of this species 
liy which it can easily be recognised, but he does not i-efer to the 
presence of a setose pad, as in C. gibbula, on the antero- ventral 
part of the abdomen. 

Distribution. Widely distributed in the Indian Ocean, Tropical 
and South Pacific. 

OORYCELLA CARINATA Giosbr. (PI. XI. fig. 10.) 

In five gatherings, 55 specimens. 

Length of females '85 mm. 

The characteristic features of this species are the cephalon (deep 
from front to back), the small eyes moderately far apart, the long 
slender thoracic spines, and the abdomen, widest at its anterior 
fourth (excluding furca), with a ventral setose pad. The furca is 
about half as long as the rest of the abdomen. 

CoRYCELLA CURTA, Sp. n. (PI. X. figS. 7-11, PI. XI. figS. 1-6.) 

Female (PI. XI. figs. 1, 2).— Length "7 mm. Very close in 
general appearance to G. carinata, bvit may be distinguished by 
the more slender build in lateral view, by the shorter thoracic 
spines, which do not reach as far as the genital openings, and by 
the absence of the patch of fine setfe or spinules on the antero- 
ventral part of the abdomen. 

The furca is short, about 2x7, and contained 2| times in the 
length of the rest of the abdomen (PI. XI. fig. 3), which is 
broadest at its anterior fourth and tapered posteriorly. 

The appendages have no special features. The fine serrulation, 
found on the third joints of the exopods in some species, is absent 
(PI. X. figs. 7, 8). The 2nd basal of the fourth foot (PI. X. fig. 11) 
is rounded on its inner face, and does not form one straight line 
with the inner face of the 1st basal. The angle between the 
base and inner margin of the 2nd basal is slightly obtuse. 

Only one specimen was found. 

In the same gathering there occurred two specimens which 
seem to be the males of the above, on account of the similarity of 
their thoracic spines, fourth feet and furca, and their small size. 

Male (PI. XI. figs. 4-6). Length '65 mm. Cephalothorax in 
middle line "4 mm., abdomen and furca '24 mm. The form of 
the abdomen (PI. XL fig. 4) is more easily shown in the figure 
than described. The outer edges of the 3rd joints of the exo- 
podites of 1st and 3rd feet are not serrulate. The angle between 
the base and inner margin of the 2nd basal of the 4th foot 
(PI. XL fig. 5) is slightly obtuse. 



FROM CHEISTMAS ISLAND. 287 

CORYCELLA Sp., S • (PL XI. figS, 8, 9.) 

There were present numerous specimens of males of CoryceUa 
belonging to at least two species judging by size alone, as in other 
characters they seemed to be in agreement. These doubtless 
represent the males of C. gibbula, C. concinna, and C. carinata, 
but I have not been able to i-efer them to their respective females. 
In size the brger measured "85 mm. and the smaller •78-*8 mm. 
Dahl's (6) recognition of Dana's C. grciciUs is based on a supposed 
identification of a figure of a male ; but Dana's (9) figures would 
apply equally well to any one of these specimens, and I do not 
think that there are any giounds for regarding his description 
as valid. 

The proportional length of abdomen and furca in these speci- 
mens was 7 : 3 ; in one of Dana's figures of 0. gracilis it is 3 : 2 and 
in the other 4 : 2. Cleve (2) has given figures of the male of 
C. gibbula, which agree fairly well with my specimens, except 
that the width of the abdomen is greater in his figures. He 
states that the propoi-tion of abdomen to furca is 2:1, but in one 
figure shows it as 9 : 5, and in the other as 8:3, so it is possible 
that he may have been dealing with more than one species. 

The figures (PI. XI. figs. 8, 9) are taken from a specimen of the 
larger form. 

Genus Coryc^us. 

Of the genus Corycceus, as restricted above, there is a 
considei'able number of more or less well described forms, which 
may be conveniently summarised under the grouping adopted 
by Dahl. 

Of the forms with a very long furca, a convenient but hetero- 
gd^neous group, Giesbrecht has figured C. longistylis and C.furcifer, 
while C. lautus of Dana is easily recognised from his figure. 

The species of which the females have a one-jointed abdomen 
are, according to Giesbrecht, C. alattis, C.flaccus, and C. elongatns ; 
to them Dahl adds C. limbatus of Brady, originally described 
from a male. 

Cm'ycceus rohustus stands alone in having the end claw of the 
2nd antenna elongated in both sexes. 

Of the forms without setae on the genital segment of the 
female, all of comparatively large size, Giesbrecht has dealt with 
C. sjieciosus, C. dance, and C. ovalis, while Dahl accepts as a 
valid description the C vitreus of Dana. 

The remaining species, which do not fall into any of the above 
groups, are mostly of very small size. They all have setfe at the 
genital pores of the female abdomen, and may be divided into 
two groups according as the endopodite of the fourth foot bears 
one or two setse. Of the first group C, obtusus, C. gracilicaudatus, 
and C. venustus ha%'e been figured by Giesbrecht ; C. minutus, 
C catus, and 0. paciiicus have been described, in his synoptic 
table, without figures, by Dahl, who also admits C. lattis of Dana 



^88 MR. G. P. FARUAN ON COPEPODA 

and C. huxleyi of Lubbock. Of the second group, with two 
setfe on the endopodite of the fourth foot, C luhhocki and 
G. tenuis have been figured by Giesbrecht ; C anglicus is a well- 
known species in British waters ; C. amazonicus, C. asiaticus, 
C. minimus, and C. africanus figure as new species in Dahl's table ; 
and C erythrceus has been described and figured by Oleve (4). 
The three new species described below belong to the last group. 

CoRYCiEUS LONGiSTYLis Dana. 

C. longistylis Dana (9). 
C. variihs Brady (1). 
C. longistylis Giesbrecht (10). 
Present in four gatherings, 5 females and 7 males. 
Both sexes of G. longistylis are easily distinguished from 
G. lau,tus and G.furcifer, which also have a long furca, by the 
broad spreading 3rd thoracic segments. The female is not com- 
pletely figured by Giesbrecht, but Brady's figure of the whole 
animal (1) (pi. Hi. fig. 1), under the name G. varius, is fairly 
aecura,te. 

Distribution. Indian Ocean, China Sea, and Tropical Pacific. 

CoRYC^us LAUTUS Dana. 

G. Icmtits Dana (6). 
G. lautus Dahl (5). 

Present in four gatherings, 12 females and 29 males. 

The female is distinguishable from that of G.ftm-cifer by its 
larger size, 2'75 mm., G. ftircifer $ measuring only 1-8 mm., 
and by the form of the abdomen, the anal segment being much 
wider in front than behind, while in G. furcifer it is only slightly 
tapered. The male of C. lautus (2* 15 mm.) is consideralbly larger 
than that of G.furcifer (1'3-1"45 mm.), but otherwise resembles 
it rather closely in general appearance. The furca, however, is 
only li times as long as the rest of the abdomen, instead of 
almost \\ times as in G. furcifer S • 

Both species can be readily distinguished by dissection, G. lautiis 
having two setse on the endopodite of the fourth foot, while iii 
G.furcifer there is only one. Dana's (9) figure of G. lautus 2 is 
easily recognisable as distinct from G.furcifer. 

Distribution. Kingsmill I. (Daoia), N.E. Atlantic. 

OoRYCiEUS ALATUS Gicsbrecht. 

The female of G. alatus is well represented in the collection, 
36 specimens occurring in five gatherings. The shape of the 
abdomen, as well figured by Giesbrecht, makes it unmistakable. 
The males of this species and G.fiaccus present some difficulties. 
While the females of G. alatus and G.fiaccus number respectively 
36 and 12, there are 76 specimens of a male which corresponds 
closely to Giesbrecht's (10) description of G. elooigatus d • These 
specimens axe undoubtedly the males of G. alatus and G. flaccus, 



FROM CHRISTMAS ISLAXD. 289 

but I have )ieeii unal)le to distinguish between them or to 
separate them satisfactorily from C. elongatas S • They can be 
readily picked out under a dissecting microscope, owing to the 
fact that, apart from differences of form, the maxillipedes are 
coloured a rusty- red. 

Distribution. Tropical Pacific. 

CoKYOiEus FLACcus Giesbrecht. 

There is nothing to add to Giesbrecht's (10) figures and de- 
sciiption of this species, which is easily separated from (J. alatus, 
tlie only species which it at all resembles, by the form of the 
abdomen and furca. There occurred twelve specimens, females, 
in three gatherings. 

The other two species of this group, G. elongatas and G. lirn- 
hatus^ which Dahl (5) regards as distinct, were not found in the 
collection. 

Distrihation. Mediterranean, Indian Ocean, Tropical Pacific. 

CoRYC'.Eus ROBUSTUS Giesbrecht. 

G. venastns Brady (1). 

G. rohustus Giesbrecht (10). 

Present in three gatherings, 10 males and 2 females. 

The female is i-ecognisable by its very lai-ge size and by the 
form of the genital segment, which is provided with lateiul ridges 
and overlaps the anal segment in dorsal view. 

There can be no doubt that the species which Brady (1) re- 
corded as G. venustus Dana was in reality G. robustios. The 
detailed figure of the abdomen shows the genital segment over- 
lapping the anal, in a way wliich is characteristic of that species 
alone. The figure (pi. liv. fig. 8) of the whole animal is 
somewhat misleading, as the artist or engraver evidently, after 
the picture was finished, mistook the lateral abdominal ridges 
for egg-sacs, and touched them up with a view to increasing the 
i-esemblance. Anyone who compares Giesbrecht's figui-e of 
G. rohiostus $ (11) (pi. li. fig. 38) with Brady's figure of G. ven- 
■astus (1) (pi. liv. fig. 8) can hardly avoid the conclusion that 
both had the same species before them. The dimensions given 
by each are identical. Brady's description does not mention any 
characters of specific value. Giesbrecht's suggestion that Brady's 
G. venustiis was really G. ohtusus seems to me to be quite i)aseless. 

Distrihation. Red .Sea, Arabian .Sea, Indian Ocean, Tropical 
Pacific, and off Cape of Good Hope. 

CoRYC.EUS SPECiosus Dana. 

An easily recognised species, well represented in tlie collection. 

It occurred in six gatherings, 22 females and 33 males. 

Distrihution. Mediterranean, Pted Sea, Arabian Sea, Indian 
Ocean, Tropical Pacific, ofl:" Cape of Good Hope, Tropical and 
North Atlantic. 



290 MR. a. p. FABRAN ON COPEPODA 

CoRYC^us DANJE Giesbrecht. 

Dahl (5) lias proposed that Giesbi-echt's name of C. danm 
should give way to Dana's G. crassmsctdus which he belieA'es to 
repi-esent the male of the same species. There is some resem- 
blance in the figure given by Dana to G. danm S , but there does 
not seem to be any reason for upsetting a well-established name 
with a recognisable description on account of its possible identity 
with a very imperfect description with insufficient figures, to 
which, at best, it can only be brought home by a process of 
exclusion. 

G. dance occurred in seven gatherings, 44 females and 70 males. 

Disiribtotion. Mediterranean Sea, Red Sea, Arabian Sea, Indian 
Ocean, Tropical Pacific, oif Cape of Good Hope. 

OoRYCJEus GRACiLicAUDATUs Giesbreclit. (PI. XI. figs. 11, 12.) 

Dahl (5) has proposed that Giesbrecht's name should be re- 
garded as a synonym of Dana's G. agilis, which i-epresents a male, 
length yLy inch or '83 mm. Females, which are almost certainly 
Giesbrecht's G. gracilicaudatus, occurred in five gatherings, nine 
specimens. In Dahl's key they fall in with G. agilis. 

What I believe to be the male of the species occuried in three 
gatherings, 50 specimen* (PI. XI. figs. 11, 12). The length was 
•76 mm. The proportions of the abdomen and furca agreed 
fairly well with Dana's figure of G. agilis, but the 3rd thoracic 
segment had the distance between its points less than the width 
of the hinder end of the 2nd thoracic segment. In Dana's figuie 
the points are spread widely outwards. These males^; wei-e 
i-emarkable in having the longest fui-cal seta, bright red. 

Distrihution. Red Sea, Arabian Sea, Indian Ocean, Tropical 
Pacific. 

CoRYC^us CATUS Dahl. (PI. XII. figs. 1-3.) 

Dahl (5) has given the above name to a species of Govycmus, 
the female of Avliich is only to be distinguished, according to his 
diagnostic table, from G. ohtusits {G. ovalis of Dahl) by having 
outstanding sharp points on the ends of the 4th thoracic segment, 
whereas in G. obtusus the ends of that segment are short and 
blunt. The other chaiacters, extracted from his table, are — 
Endopodite of 4th foot with one seta ; abdomen with seta behind 
the genital opening ; size not more than 1 mm. ; furca compara- 
tively short ; third thoi-acic segment spreading, more than | as 
broad as fii-st, the points not reaching to the distal end of the 
genital segment ; anal segment very little longei' than its basal 
width. 

There are several specimens of a female Govi/cceus in the 
collection which agree with the above diagnosis and aie eithei' 
G. catus or a new species, the probability being in favoui- of the 
former view. They occur in five gatherings, 90 specimens in 
all. The 3rd thoracic segment in doisal view, though spreading. 



FROM CHRISTMAS ISLAND. 291 

is not so wide as in 0. obiiosas, being at its widest pai't, a little 
Ijehind the anterior margin, only just ^ as wide as the widest part 
of the cephalon (PI. XII. fig. 1). The points of the segment are 
slightly conti'acted, the distance between them being only | of the 
width of the cephalon. The total length is '9 mm. 

The most noticeable features, in comparison with C. obkosics, 
are the sharp slender points to both the 3rd and 4th thoracic 
segments (PI. XII. fig. 2), and the somewhat larger eyes, separated 
by I'ather less than theii- own dianietei-. The foi-m of the 
abdomen and furca is v'eiy similar in both species. 

CoRYc^us OBTUsus Dana. (PI. XII. figs. 4-6.) 

Cor>/cc8i(,s obtasas Dana. 
C. obtusus Giesbrecht. 
0. ooalis Dahl. 

According to Dahl (5), Giesbrecht is in error in his identification 
of the species which he has described and figured under the 
name of G. obtasus Dana, Dana's species being different and not 
found sxibsequently. He believes that Giesbrecht's description 
really refers to the species which Glaus (2) had described under 
the name of C. ovalis. 

Under these circumstances the course least liable to cause 
confusion would be to reject both Dana's and Glaus's names ; 
however, for the present, I assume that Giesbrecht's name is 
correct. Eighteen specimens, females, of the species in question 
occurred in the collection, in three gatheiings. 

The distinguishing characters are the size, 1'05-1"1 mm., the 
wide spreading third thoracic segments, the bhmt tips to the 
fourth thoracic segments (PI, XII. fig. 5), the fine seta^ in front 
of the genital openings, the anal segment as long as it is bi'oad 
at the base and nearly as long as the furca, and the single 
seta on the endopodite of the fourth foot (PI. XII. fig. 4). 

Distribution. Red Sea, Arabian tSea, Indian Ocean, Tropical 
Pacific, Tropical Atlantic. 

CoRYC^us TENUIS Giesbrecht. (PI. XII. figs. 8, 9.) 

Of the forms with tv/o seta) on the endopodite of the foui'th 
foot and a moderately long furca there are two species, females, 
in the collection. One of these is described below as ne^v, the 
other appears to be ('. tenuis. In Dahl's (5) key the choice lies 
between C. anglic>(,s and C. tenuis, the former being distinguished 
in the key by the presence of a corner on the inner side of the 
lateral prolongation of the 3rd thoracic segment and by a small 
hook-like ventral pi-ocess on the j^roximal end of the abdomen. 
Both these characters are present in my specimens and, if the 
table were closely followed, they ought to be C. anglicihs. They 
ai-e, however, not that species, being distinguished fi-om it Ijy the 
shorter genital segment with much more slendei- seta?, the form of 
the anal segment, which is parallel-sided and about twice as long 



292 MR. G. p. FAHRAN ON COPEPODA 

as wide, and the fuvca, which is slightly longei' and more slender. 
They agree well, on the other hand, as far as the text and figures 
go, with Giesbrecht's description of G. tenuis, though that writer 
makes no mention of the ventral hook on the genital segment. 

The length, r05 mm., is slightly greater than that given by 
Giesbrecht, viz. "9 mm. The length of the cephalothorax in the 
middle line, dorsal view, is '64 mm., of the abdomen and furca 
•41 mm., the proportional lengths of abdominal segments and 
furca being 14 : 8 : 18. Measured along the ventral margin the 
proportions are more nearly 12 : 10 : 17. The pigment of the 
eye-apparatus shows very distinctly thiough the body as of a blue- 
green colour. 

The second antenna (PI. XII. fig. 9) bears a long spine on the 
first basal joint and a much shorter spine, one-third as long, on 
the second basal joint. The inner margin of the second basal ends 
distally in two strong teeth. The terminal claw of the second 
antenna is just equal to the spine of the second basal. 

The swimming-feet have no distinctive features, except that the 
terminal spine of the exopodite of the second foot is very slightly 
cui^ved. 

Six specimens were found in two gatherings. 

Distribution. Indian Ocean and Tropical Pacific. 



CoRYC^us DUBius, sp. n. (PI. XII. fig. 7, PL XIV. figs. 5-9.) 

In the Maldive collections Dr. Wolfenden (11) obtained a 
specimen of Gorycmus, which, following Dahl's table, ought to be 
C. amazonicas, but, as he remarks, without figures it is impossible 
to refer it with ceitainty to that species. In the Christmas 
Island collection there occurred one specimen of a female, 
described below under the name of C. duhius, which may well be 
the same species as that to Avhich Dr. Wolfenden refers, but 
again the absence of figures precludes certainty. The particulais 
given by Dr. Wolfenden — namely, the size, 1*0 mm., and the 
proportionate lengths of abdomen and furca 7:8: 9 — come very 
near '97 mm. and 21 : 23 : 28, the corresponding measurements 
in my specimen. The lateral prolongations of the third and 
fourth thoracic segments are short in both specimens. In 
Wolfenden's specimen, however, the spine on the second joint of the 
second antenna is said to be " not more than half the length of 
that of the 1st basal, and shorter than the distal hook," but in 
mine the spine in question is just one-third as long as that on 
the 1st basal and is distinctly longer than the distal hook. 

Cleve (4) has described a species, C. erythrceihs § , from the 
Red 8ea, Avhich is very closely allied to both the above-mentioned 
forms, but if his description be taken as accurate it must be 
distinct. The female genital segment has seta? on the genital 
openings and a ventral hook, the endopodite of the 4th foot has 
two setce, and the furca is moderately long. The size -88- '94 mm. 
is slightly less than that of G. cliMas. The proportions of the 



FKOM CUKLSTMAS ISLAND. 293 

abdominal segments and fiirca are 10 : 10 : 12, «. e, the genital 
and anal segments are equal ; in the figure, however, the genital 
segment is shown as being the longer. The most noticeable 
difference is in the 2nd antenna, in which the spine of the 2nd 
basal is figured as being half as long as that of the 1st basal, and 
is said to reach to the distal margin of the joint. In my species 
the spine of the 2nd basal is only one-third as long as that of the 
1st, and falls short of the distal margin of the joint by half its 
own length. 

In view of this uncertainty it seems advisable to give the 
Christmas Island specimen a distinct name until the specific 
chaiucters of C. amazonicus aiid C. erijthrcei(,s are more fully 
known. 

Female (PI. XIV. fig. 5).— Length -97 mm. ; cephalothorax in 
mid-dorsal line -58 mm. ; abdomen -39 mm. (PI, XIV. fig. 6) ; 
projiortions of abdominal segments and furca, dorsally 21 : 23 : 28, 
ventrally 16:22:27. Sides of cephalon parallel, Ceph. + Th. 1 
being about twice as long as broad. Points of 3rd thoracic seg- 
ment (PI. XIV. fig. 7) not very long (broken in my specimen), 
with thickened margin on posterior edge. 4th thoracic segment 
short, with sharp points. Genital segment about 1 j times as long 
as wide, with setse on genital openings, in lateral view with a 
small sharp hook anteriorly on ventral edge. Anal segment a 
little longer than genital segment and slightly overlapped by it 
dorsally, about three times as long as wide. Fui^ca slightly 
diverging, a little longer than anal segment. Furcal rami about 
1x10, tei'minal setfe missing. 

1st antenna six- join ted, proportional lengths of joints } ^ ^ "^ ^ ^ . 
2nd antenna (PI. XIV. fig. 8) with spine on 1st basal very long ; 
spine on 2nd basal short, about § as long as that on 1st basal and 
falling short of the nearest point of distal margin of joint by 
about half its own length. Distal margin of 2nd basal with two 
sharp teeth. Terminal spine of 2nd antenna very strong, shorter 
than spine on 2nd basal. 

Mouth-parts not satisfactorily made out. 
Jointing and number of setee of swimming feet normal. 
1st foot : exopodite with broad terminal spine, almost as lono- as 
endopodite, with fine denticulations. 

2ud foot (PI. XIV. fig. 9) : terminal spine of exopodite almost 
straight, with moderately coarse denticulations on outer edge and 
two small distal serrations on inner edge (possibly an individual 
eharacter and not specific). 

3rd foot with long slender terminal spine on exopodite, the 
proportional lengths of joints and spine being 28 : 20 : 60 : 70. 

4th foot (PI. XII. fig, 7) : endopodite small, with two setje ; 
exopodite slender, the proportional lengths of its joints beino- 
approximately 6:4:5; outer edge seta on 1st joint almost 
as long as 2nd joint, blunt process on 2nd basal joint internal to 
endopodite. 

Proc. ZooL. See— 1911, Xo. XX. 20 



294 MR. G. p. FARRAN ON COPEPODA 

CORYC^EUS ANDREWSI, Sp. 11, (PI. XIII. figS. 7-9, PI. XIY. 

figs. 1-4.) 

Female (PI. XIV. fig. 1). — Length -95 mm. ; ceplialothorax in 
middle line "64 mm., to end of thoracic spines -Tmm. ; abdomen 
•31 mm. (PI. XIY. fig. 4). Proportional lengths of abdominal 
segments and furca 19 : 12 : 10. F'urcal rami about five times as 
long as wide. The genital segment is about | as wide as long 
and 4 as deep, in lateral view, as long ; the antero-ventral angle 
is very marked in lateral view, being almost a right angle, 
rounded off at the apex and furnished with a small patch of 
minute spinules. The points of the 2nd . thoracic segment are 
short and notched on the inner margin. The points of the 4th 
thoracic segment are very short, but sharp at the extreme tip. 

The eyes are of medium size and are separated by rather less 
than their own diameter. 

1st antenna, proportional lengths of joints ^ 3 ^ ^ 3 2 - 
2nd antenna (PL XIIL. fig. 9) : 1st basal with long spine ; 2nd 
basal with spine about | as long as that on the first, and two 
rather slender teeth on its distal edge. The terminal spine of the 
2nd antenna is about half as long as that on the 2nd basal. 

The pi-oportional lengths of the joints of the swimming-feet are 
best seen in the figures. 

The outer edge spines of the exopodites of the 1st and 2nd feet 
(PI. XIII. fig. 8, PI. XIY. fig. 3) have well-developed laminaj, but 
on the 3rd foot they are very slightly lamina,te. The terminal 
spine on the exopodite of the 2nd foot is slightly curved, on the 
1st and 3rd feet it is straight. In the 4th foot (PI. XIII. fig. 7) 
the outer edge si3ine of the 1st joint of the exopodite is short and 
slender. The endopodite of the 4th foot bears two moderately 
large setse. 

Five specimens, females, were foiuid in two gatherings. 
Comparing G. anch-ewsi with the other species in which there 
are two setse on the endopodite of the 4th foot, we find that it is 
separated from C laiUus, C. luhhocki, C. tenuis, G. erythrcetis, 
G. venustus, G. anglictts, and G. africanus by its short furca, 
which is less than ^ of the length of the rest of the abdomen. 
In G. amazonicus the anal segment is said to be longer below 
than the genital, a fact which at once distinguishes tha,t species. 
In G. asiaticits and G. 'minimus, according to Dahl's table, the 
furca agrees in being about half as long as the genital segment, 
but the genital segment is snid to be respectively two and three 
times as long as the anal, while in G. andreivsi it is only 1| times 
as long. 

CORYC^US MURRAYI, Sp. n. (PI. XIII. figS. 1-6.) 

Female (Ph XIII. figs. 1, 2). — Length 1*2 mm,,: eephalothorax 
in middle line '75 mm. ; abdomen and furca '43 mm. The pro- 
portional lengths of the abdominal segments and furca in "01 mm. 
are 18 : 13 : 12. The wings of the 3rd thoracic segment 



FROM CURISTMAS ISLAND. 295 

(PI. XIII. fig. 3) fire long, sharp-pointed, and spi^eading, the 
width between the points being § of the width of the cephalon 
and equal to the length of the abdomen without furca. The 
.abdomen is wide in front and regularly tapered to the furca, 
the segmentation between the genital and anal segments being 
ratlier obscure. Immediately behind and exterior to each genital 
opening is a small blunt process or papilla projecting beyond 
the margin of the genital segment in dorsal view. The setfe 
at the genital openings are very small and might easily be 
overlooked. 

The 2nd antenna (PL XIII. fig. 4) is much stouter than in 
C. andreiosi and has one strong tooth on the inner margin of the 
2nd joint. The seta on the 2nd joint is rather more than half as 
long as that on the 1st joint, and reaches almost to the end of 
the inner edge tooth. The terminal claw of the 2nd antenna is 
strong. 

The swimming-feet have no special characters. The terminal 
spine of the exopodite of the 2nd foot (PI. XIII. fig. 6) is only very 
slightly curved. The endopodite of the 4th foot (PI. XIII. fig. 5) 
bears two moderately strong setae, and the outer edge spine of the 
1st joint of the exopodite is i-a.ther shoi^t. 

This species is rather closely allied to the above-described 
C. andreiosi, having, like it, setae on the female genital segment, 
two setfe on the endopodite of the fourth foot, and a short fvirca. 

Six specimens were found in three gatherings. 



Literature referred to. 

(1) Brady, G. S. — 'Challenger' Reports^ vol. viii. pt. xxiii. 

Copepoda, 1883. 

(2) Claus, C. — Die freilebenden Copepoden, 1863. 

(3) Cleve, p. T. — " Plankton from the Indian Ocean and 

Malay Archipelago." Konigl. Sv. Vet.-Akad. Hand!., 
Bd. 35, No. 5, 1901. 

(4) Cleve, P. T.— " Plankton collected by Mr. Thorild Wulff' 

during a Voyage to and fi'om Bombay." Arkiv fiir 
Zoologi, Bd. i., 1903. 

(5) Dahl, F.— Verb. Deutsch. Zool. Ges. 1894. 

(6) Dana, J. D. — Proc. Acad. Nat. Sci. Philadelpliia, ii , 

1845. 

(7) Dana, J. D.— Ann. Nat. Hist. vol. xviii., 1846. 

(8) Dana, J. D. — Proc. Amer. Acad. Sci. ii., 1849. 

(9) Dana, J. D, — U.S. Explor, Exped., vol. xiii. Crustacea, 

1852. 

(10) GiESBRECHT, W. — Fauna u. Flora des Golfes von Xeapel, 

vol. xix. Copepoda, 1893. 

(11) WoLFENDEN, R. N. — Fauna and Geography of the Maldive 

and Laccadive Archipelagoes, vol. ii., suppl. 1, Copepoda, 
1905. 

20* 



296 ON COPEPODA FROM OURISTMAS ISLAND. 

EXPLANATION OF THE PLATES. 

All the figures have been drawn with the aid of a camera lucida. 

Plate X. 
Fig. 1. CoryceUa hrcvis $ , dorsal view. 



2. 






„ 9 , lateral view. 


3. 


jj 




„ ? , 3rd foot. 


4. 


jj 




„ $ , 1st foot. 


5. 


jj 




? , 4th foot. 


6. 


)3 




„ $ , 3rd thoracic segment, mounted 


7. 


Coryc 


ella 


curta ^ , 3rd foot, exopodite. 


8. 


jj 




„ ?,2iidfoot. 


9. 


jj 




„ ? , 3rd thoracic segment, mounted 


10. 


jj 




., $ , 2nd antenna. 


11. 


„ 




„ '^ , 4th foot. 



PiATE XL 

Fig. 1. Cori/celJa curta $, dorsal view. 

2. „ „ ^ , lateral view. 

3. „ „ $ , abdomen, dorsal view. 

4. „ „ (^, abdomen, dorsal view. 

5. „ „ ^, 4th foot. 

6. „ ,, (?, 3rd thoracic segment, mounted. 

7. Corycella brevis $ , 2nd antenna. 

8. C'oryceZZa sp., (?, dorsal view. 

9. „ „ (? , 4th foot. 

10. Corycella carinata $ , abdomen, lateral view. 

11. Corycceiis gracilicaudatus (3' , dorsal view. 

12. ,, „ (J , lateral view. 

Plate XIL 

Fig. 1. CoryccBus catus ? , dorsal view. 

2. „ „ $ , 3rd and 4th thoracic segments, mounted. 

3. „ „ ?, 4th foot. 

4. Corycceiis obtusus $ , 4th foot. 

5. „ „ $, 3rd and 4th thoracic segments, mounted. 

6. „ „ $ , 2nd antenna. 

7. Cmycceus dubius ^ , 4th foot. 

8. CoryccBus tenuis $ , 3rd and 4th thoracic segments, mounted 

9. j, „ $ , 2nd antenna. 

Plate XIII. 



Fig 


. 1. 


Coryc(Bus murrai/i 


9 , dorsal view. 




2. 


» » 


9 , lateral view. 




3. 


» )! 


^ , 3rd and 4th thoracic segments, mounted 




4. 


)) 


!^, 2nd antenna. 




5. 




? , 4th foot. 




6. 


7) 


? , 2nd foot. 




7. 


Corycceus andrewsi 


$ , 4th foot. 




8. 


» „ 


? , 2nd foot. 




9. 


„ „ 


^ , 2nd antenna. 



Plate XIV. 

Fig. 1. CoryecBus andrexosi 9 , dorsal view. 

2. ,, ,. $ , 3rd and 4th thoracic segment.s, mounted. 

3. „ „ ?, 1st foot. 

4. ., ,, $ , abdomen, dorsal view. 

5. Corycaus dubius 2 , dorsal view. 

6. „ » ? 7 abdomen, lateral view. 

7. „ ;; ? ) 3rd and 4th thoracic segments, mounted. 

8. „ „ ? , 2nd antenna. 

9. ,, ,; ? ! 2nd foot, exopodite. 



ox SOME NEW ZEALAND Sl'll>Eli.S. 2U7 

14. On some Now Zealand Spiders. 
ByH. R. Hogg, M.A., F.Z.S. 

[Received November 11, 1910: Head February 7, 1911. J 

(Text-figures 92-96.) 

Prof. Chas. Chilton, of Christchurch, New Zealand, suggested 
to his students that they should in the vacation collect specimens 
at various points over the Noi'th and South Islands, wherever 
their homes were situated. The result is that from some half- 
dozen localities specimens of spiders were brought in within the 
space of a few weeks, and these he has kindly sent to me. 

New Zealand for a considerable period has been rather well 
worked from a zoological point of view, and its spider fauna 
described not only by Dr. Koch but by local men such as 
Messrs. Urquhart and Goyen. It is therefore not a little re- 
markable to find in the small collection here described so large 
a pi'oportion of new species, and, moreover, the same new species, 
in one case from three, and another from two widely separated 
localities at the same time. The thirty specimens comprise no 
less than twelve species of eleven genera. Of the species, four 
are new, and one a local variety of an Australian species not 
hitherto recorded from New Zealand. They were collected mostly 
from Ruakura and Wellington, in the North Island, by Miss B. 
D. Cross, and one by Mr. E. P. Turner from the summit of 
Mt. Ngauruhoe shortly after an eruption. The latter appears 
to have minute particles of volcanic dust adhering to it. 

The species from the South Island were taken by Messi's. 
R. N. Hawkes and G. M. Thomson, and by Prof . Chilton himself . 

On Mt. Peel, in the Province of Canterbury, Mr. Hawkes 
picked up at random a male Uliodon of a new species, a new 
species of Ar-gocteuus, a male of Cainhridgea.antipodiana (with 
its unique form of stridulatory organ), and three females of For- 
rhothele antiiJodiana Walck., a new locality for the two latter 
species. 

In 1905 (Zool. Jahrb. xxi. pt. 4, 1905) M. Simon formed a genus 
Mynoylenes for a spider (J/, iiisolens) from the Chatham Islands, 
500 miles east of New Zealand, a,nd I was recently able to describe 
another species (Reports Phil. Inst, of Canterbury, N. Z., 1909, 
vol. i. p. 165) from the Sub- Antarctic Auckland and Campbell 
Islands, south of New Zealand, but the genus had not so far been 
recorded from New Zealand itself. Miss Cross, Prof. Chilton, and 
Mr. Turner send specimens, one each from separate localities: — 
Ruakura and Mt. Ngauruhoe in the North, and Picton m tiie 
South Island. These, although differing in size, are all adult and 
appear to be similar. I have taken them to be the same species. 

Hemicloea rogenJioferi L. Koch and Araneus verrucosus Walck., 
from Ruakin^a, are among the few spiders hitherto known to 
be common to both New Zealand and Australia. ; and Tetragnatha 



298 MR. H. R. HOGG ON SOME 

ferox L. Kocli, with some slight variations, is now for the first 
time recoi'ded from New Zealand. 

Prof. Chilton sends Desis marina Oamb., taken by himself in 
Lyttelton Harbour, and a Pardosa from Picton Harbour. 

Suborder MYGALOMOKPH^. 

Family AYICULARIDJE. 

Subfamily Diplurin^e. 

Group Machotheleje. 

Genus PoRRHOTHELE Simon. 

PORRHOTHEtE ANTIPODIANA Walck. 

Three females from Mt. Peel, Canterbury. Collected by 
Mr. R. N. Hawkes, 

Suborder AEANE^ VEE^. Section ECRIBELLAT^. 

Family DRASSID^. 

Subfamily HEMiCL(EiNiE. 

Genus Hemiclcea Thor, 

Hemiclcea rogenhoferi L, Koch, 

One male from Ruakura. Miss B. D. Cross. 

Family ARGIOPID^. 

Subfamily Argiopin^. 

Group Tethagnatheje, 

Genus Tj^tragnatha Latr, 

Tetragnatha ferox L. Koch, var, cross^e, nov. 

One male and two females from Ruakura. Miss B. D, Cross. 

A few variations from the Australian form are noted below, 
but it is so like in other respects that I do not think it can be 
specifically separated therefrom. 

Group Arane.^^, 
Genus Meta Clerck. 
Meta argentata Urquhart. 

Nephila argentatum Urq. Trans. N.Z. Inst. vol. xix. p. 92. 
One male and two females from Ruakura, Miss B. D. Cross. 
These correspond so nearly in all respects with Mr, Urquhart's 
description of the i\'^e^7AiZ« above mentioned, except that the side eyes 



NEW ZEALAND SPIDERS. 299 

are close together, that I have no doubt ns to theiv belonging to 
tlie same species. The silvery colouring of the abdomen is 
suggestive of some of the Nephilcti, especially N. venosa L. Koch, 
but the shape of its cephalothorax and shorter lengths of tarsal 
joints show that it is not a Nepliila^ of which genus no species 
liave been recorded from New Zealand. 

Genus AuANEUs Linn. 
Araneus verrucosus Walck. 
Two males and six females from Ruakura. Miss B. 1). Cross. 

Family CLUBIOKID^ 

Subfamily C t e n i n .e. 

Group Ctene^'E. 

Genus Uliodon L. Koch. 

UlIODON HAWKESl, Sp. n. 

One male and one female fi-om Mt. Peel, Canterbury. 
Mr. R. N. Hawkes. 

One male fi-om Wellington, Miss B, D. Cross. 

Subfamily Liocranin^. 

Group MlTUIlGE-E. 

Genus Argoctenus L. Koch. 
Argoctenus aureus, sp. n. 
One female from Mt. Peel. Mr. R. N. Hawkes. 

Family AGALENID^. 

Subfamily C y b .e i N js. 

Group Besiege. 

Genus Desis Walck. 
Desis marina Cambr. 
One female from Lyttelton Harbour. Prof. Chilton. 

Gi'oup Cyb^ee,e. 
Genus Mynoglenes Simon. 
Mynoglenes chtltoni, sp. n. 

One male from Ruakura. Miss B. D. Cross. 
One male from Mt. Ngauruhoe, Waikato, IST. Island. Mr. E. P. 
Turner. Shortly after an eruption. 

One female from Picton, S. Island. Prof. Chilton. 



300 MR. H. R. HOGG ON SOME 

Group Abgyhonete^^:. 
Genus Oambridgea. 
Cambridgba antipodiana White- 
One female from Mt. Peel, Mr. R. N. Hftvvke.s. 

Family PISAURID^E. 

Group DOLOMEDEM, 

Genus DoLOMBDES Latr, 

DoLDMEDEg TRIDENT ATUS, Sp. n, 

One female from Dunedin, Mr, G. M. Thomson. 

Family LYCOSID^, 

Group Pardose^. 

Genus Pardosa C, Koch, 

(Probably) P, cajjescens Goyen, 

Lycosa canescens Goyen, Trans. JST.Z, Inst, vol, xix. p. 203. 

The specimens, one male and one female, agree closely with 
Mr, Goyen's description of his species, which he states he always 
found on river banks. Prof, Chilton collected this from a mud- 
bank in Picton Harbour, and I have little doubt as to its being 
the same. Its short broad lip, square front, two large equal 
teeth and one smaller on the inner margin of the falx-sheath, and 
procurved front row of eyes, show clearly that it belongs to 
0. Koch's genus Pardosa, and this would follow from Mr. Goyen's 
description of the lip alone, 



Tetragnatha PER0J5: L, Koch, var, crosSjE, nov, (Text-fig. 92.) 

Female, Cephalothorax, mandibles, maxillae, coxfe, and legs 
orange-yellow. The lip, except on the front edge, is yellow-brown 
and the stejrnum somewhat darker. The scanty hairing is rather 
pale brown, and the spines on the legs dark brown. The cephalic 
part of the cephalothorax is bordered with a somewhat browner 
line, but with white hairs thereon, and there are two jDairs of 
rather darker spots on the front of the rear slope. The abdomen 
above is yellow-grey, bordered with a dark grey wavy line on each 
side, the whole distance from front to rear. There are short white 
hairs rather thicker on the side slopes than on the back. Under- 
neath it is a dingy yellow-grey with short brown hairs; there is a 
rather darker area running from the spinnerets to the breathing- 
apertures and epigyne. 

The cephalothorax is a long oval, not quite twice as long as 
bi"oad, rounded at the sides, the cephalic part distinctly raised above 
the thoracic. Between the end of the former apd the rear slope 



NEW ZEALAND SPIDERS. 

Text-fig. 92. 
b 



301 




Tetragnatliafero.v, vai-. crosses. Male. 

a, eyes ; b, mandible of male from inside. 

c, „ female from inside. 

d, „ male from outside. 
c, lip and maxillfE; /, epigyne; </, male palp. 



302 MR. U. R. HOGG ON SOME 

are two pairs of depressions, one behind the other, divi(.led by a 
rather broad ridge. 

Viewed from above both rows of eyes are recurved, those of the 
rear row equal and equidistant two and a half times their diameter 
apart. The front median eyes are nearer together than the I'eai', 
and apparently rather larger ; they are situated on the sides of a 
protuberance, and are as far from the rear eyes as from one 
another. The respective pairs of side eyes are a similar distance 
apart. As they are all dark and on black rims, the real size of 
the eye is difficult to distinguish. The clypeus is as broad as a 
front median eye. 

The 'mandibles are as long as the cephalothorax, divergent, and 
protrude at an angle of 45 degrees. They have long upstanding- 
hairs on the inner side and shorter ones on the outer edge. On the 
outer margin of the falx-sheath there are five rather long teeth, 
the upper one on the top edge being the largest. On the inner 
margin there is one large tooth at the top, one at the corner, then 
one small one a little way down, and then seven nearer together 
reaching to the top of the lip. On the inner side of the fang itself, 
near the base, is a short stout spine midway between the two 
marginal spines on the top edge of the sheath. 

The maxillce are long, broadest at the anterior end, where they 
rather bend back with a rounded corner at the outer edge ; from 
here they slope in to about the middle and then out again. The 
trochanter of the palp is very long. 

The lip is about two-fifths the height of the maxillse, rounded 
in front with a rather prominent edge. It broadens out towards 
the posterior end, which is also rounded, and there is a depression 
all the way round a little distance from the margin. 

The sternum is a long narrow shield-shape, twice as long as 
broad, hollowed in front, and with a prominent apophysis at each 
of the front corners ; it is smooth and shiny, with a few fine up- 
standing hairs. There is a space between the second and third 
pair of coxee, and the fourth pair are not quite contiguous to 
one another. 

The abdomen is long and narrow, straight at the sides and 
slightly rounded at the front and rear. 

The epigyne is a transverse oval, the inside filled with an oval 
convexity. 

The ^«Zpt are thin. The tibial joint is twice as long as the 
patellar. 

The legs are long and thin ; the tibia of the front pair more than 
six times as long as the patella. There are four pairs of spines at 
the sides of tibia? 3 and 4 ; the spines are fine and not very long. 

The male is coloured the same as the female, except that the 
sternum is of the same bright yellow colour as the cephalothorax, 
maxillae, and legs. 

The mandibles are quite as long as the cephalothorax ; on the 
inner margin of the falx-sheath are twelve teeth, the second of 
these being the largest. On the outer side there are three large 



NEW ZEALAND SPIDERS. 



303 



protuberances at the end, and five rather large teeth at intervals 
down the mai^gin of the falx-sheath. 

The male differs from T. ferox L. Koch in the larger second 
tooth from top on the inside of the mandible, in the apophyses at 
the head, the two inner curving in the opposite direction to the 
outer, in the mandibles being quite as long as the cephalothorax, 
instead of shorter, in the tibial joint of the palp being distinctly 
longer than the patellar, and the palpal stigma being not more 
than half as long as in T. ferox, and the legs longer in proportion. 

The measurements (in millimetres) are as follows : — 

Male, 

Long. Broad. 

Cephalothorax ... 3 2 

Abdomen 5^ 1 

Mandibles 3 

Troclianter Patella Metatarsus 
Coxa. & femur. & tibia. & tarsus. 

Legs 1. 1 9 10 11 = 31 

2. 1 7 7 7 = 22 

3. 1 ^ n 3 = 10 

4. 1 6i 6| 6i = 20A 
Palpi ,. A 21 2~ l| = 6| 

Female. 
Long. Broad. 
Cephalothorax ... 3 1| 

Abdomen 7 2 

Mandibles 3 

Pat. & Metat. 
Coxa. Tr. & fern. tib. & tars. 

Legs 1. 1 7 8 9 = 25 

2. 1 6 6 6 = 19 

3. 1 21 2 2| = 8 

4. 1 5 5 51 = 161 
Palpi k 21 2 11 = 61 

T. ferox L. Koch, fairly common along the eastern and southern 
coasts of Australia, has not been previously recorded from New 
Zealand. The New Zealand and Australian females are practi- 
cally identical, and I consider this to be a local variety, which I 
have named after Miss Cross, who sends a male and a female from 
Ruakura. 

Uliodon hawkesi, sp. n. (Text-fig. 93.) 

Female. The cephalothorax is bright, rather pale yellow-brown, 
with short upstanding bi'own bristles and downlying white haiis. 
On the cephalic part there are three longitudinal dai-ker lines — one 
median reaching from the eyes to the rear end, and one shorter 
each side, also a darker line round its margin. At the sides of the 
thoracic part is a darker brown mai-ginal stripe. 



304 



MK. H. R. HOGG ON SOME 



Mandibles davk yellow-brown, with downlying white hairs and 
upstanding brown bristles ; fangs bright red. Lip and maxilla^ 
yellow-brown, with greyish-yellow hairs. Sternum yellow-brown, 
with whitish hairs. Ooxfe pale yellow, with a dark band at the end. 
Femora yellow, with three dark bands underneath. The other 
joints of the legs dark yellow-brown, with black spines ; grey 
scopul^ on all tarsi and metatarsi. 

Abdomen above greyish -yellow mottled with black, sparsely 
covered with brown upstanding bristles and downlying white hairs. 
On the under side the fore part is yellow, with short 3fellow-brown 
hairs as far as the pulmonary apertures, the remainder pale yel- 
lowish grey. The sides are darker, nearly black, with longitudinal 
lines of yellow and white hairs. 

Text-fig. 93. 



OoqO^ 




a, e3'es ; 



JJUodon liaivTcesi. Female. 
b, male palp; c, lip and maxillae; d, epigyne. 



The ceplialotliorax is convex, highest at the beginning of the 
rear slope, thence downwards to the eye-space. The cephalic part 
is rather more than half the greatest width of the thoracic, its sides 
rather straight, but those of the thoracic rounded. Along deep 
longitudinal fovea reaches to the top of the rear slope, which is 
steep. 

The rear row of eyes viewed from above is recurved, but 
procurved from in front ; the median eyes their diameter apart 
and rather more than three times that distance from the laterals, 
which are of the same diameter. Of the shorter front row, the 
laterals are as large as those of the rear row, but the median are 



NEW ZEALAND SPTUEliS. 305 

two-thirds their diameter. These are all e({iiidistant, the diameter 
of the median apart, and one and a half of the same from the 
corresponding eyes of the rear row. 

The dyjjeus is the width of the front median eyes to the margin 
of the behaired part, and there is an equal distance of smooth 
muscle to the insertion of the mandibles. 

The mandibles are much kneed at the base, narrowing to the 
anterior end, thickly covered with bristly hairs. The fangs are lono- 
and powerful. On the inner margin of the falx-sheath are three 
large equal teeth, and one large between two smaller on the outer. 

The lip is longer than broad, hollow in front, I'ounded at the 
sides, and constricted at the basal part, which is rather long. It is 
more than half the length of the maxillae, which are narrow at the 
base and gradually wider to the upper margin, where they are 
broadest, curving gradually on both inner and outer sides. 

The sternum is a broad shield-shape, slightly convex, pointed at 
the rear between the nearly touching coxfe. It is thicklv covered 
with upstanding bristly hairs. 

The legs are stout and moderately long. The trochanters have 
a median ring of bristles rising from a chitinous ridge. The 
tibiae and metatarsi have long powerful spines, with a bunch 
at the anterior end of the latter. There are thick scopulfe on the 
tarsus and metatarsus of all legs. The two claws have five or six 
pectinations. 

The paljy of the female has a pair of long spines at the base of 
the distal joint ; the claw is smooth. 

The abdomen is ovate, pointed at rear end, and the spinnerets 
terminal. The inferior pair are conical, two-jointed, the second 
joint being short and hemispherical. The superior pair are 
cylindrical, half the length of the inferior, with similar second 
joints. 

The measurements (in millimetres) are as follows : — 



Female. 
Long. 

Cephalothorax . 



Long. Broad. 

3 in front. 

H 

Abdomen 11 7| 

Mandibles 3| 

Pat. & Metat. 
Coxa. Tr. & fem. tib. & tars. 

Legs 1. 2 6| 7 6* = 22 

2. 2 51 61 6| = 201 

3. 2 6 51 6~ = 191 

4. 2 61 f 81 = 24' 
Palpi 1 31 2 21 = 9 

One male and one female collected by Mr. R. N". Hawkes on 
Mt. Peel, Canterbury, at an altitude of 2000 feet. 

One male from Wellington, collected by Miss B. D. Cross. 



306 



MR. 11. R. nOGG OK SOME 



Besides being twice as large, this species differs from Uliodon 
{Zora) frenatttm L. Koch, from the same province, in the longi- 
tudinal lines on the cephalothorax. It has no pattern on the 
abdomen, and the sternum is yellowish instead of black-brown. 
There is a scopula on the tarsus and metatai-sus of all legs, 
instead of only on the front two pairs. 

Argoctenus aureus, sp. n. (Text-fig. 94.) 

Female. The cephalothorax is bright yellow in the middle 
the whole distance from the eyes to the rear; this is bordered on 
the side slope with dark grey, almost black, reticulations, with 
parallel stripes of the same colour reaching to the mai'gin. At 

Text -fig. 94. . 







Argoctenus aureus. Female. X 2. 
a, ej-es ; h, epigj'ne ; c, lip and maxillae. 

the margin itself, on the rear half, is a narrow band of yellow, 
and at the side of the eye-space is rather darker yellow. The 
whole is covered with short downlying white flat hairs, the rear 
fovea being black-haired. Behind the eye-space is a jet-black 
curved line passing on each side along the margin, and there are 



XEW ZEALAND SriDERS. 307 

two black patches on the clypens, but the same thick white 
hairs. Tlie mamlibles are deeper yellow, with a black streak on 
the inner edge of each, reaching from the base almost to the point, 
where it is again yellow. The fangs are red-brown. The lip is 
grey, the maxillaj pale yellow, with broad nearly white margins 
on the inner side of the former and front of the latter. The 
sternum is dark yellow-brown, thickly covered with white hairs 
and grey bristles. The legs are pale yellow on the upper side, 
with a grey streak nearly the whole length of the under side on 
the femoral, patellar, and tibial joints ; covered with white or pale 
greyish-yellow hairs. The scopulse and claw-tufts are pale grey. 

The abdomen is yellow-grey, with a brown median scollop&d 
stripe, darkened at the edges, reaching from the base to the 
spinnerets. The whole rear part is rather darker with a reticu- 
lation of grey on the yellow, the sti'ipe remaining darker still. 
The hairs are white, with the exception of a few upstanding 
bi-own ones on the stripe and at the base. The under side is 
greyish yellow, covered with white intermixed with short brown 
haii'S. The hairs on the yellow sj^innerets are brown. 

The cephalothorax is convex, rounded at the sides and broadest 
a little in front of the rear end. The front part is about one-half 
the greatest width, and the sides of this, as far back as the rear 
row of eyes, are straight. 

The eyes are in four rows. The second, row, the laterals of the 
front row, being vei'tically just clear of the median first pair. 
These are all of the same diameter and half their width apart. 
The rear two pairs are again of equal size, one-third larger than 
the front, the median ones being one-third of their diameter apart 
and the same distance from the front laterals. The median are 
14 times their diameter from the rear laterals respectively, and 
the latter are four times their diameter apai't. 

The dypeus is as broad as the front eyes. 

The mandibles are kneed at the base, divergent, and at the 
anterior end half their width at the base.* They are as long as the 
front of the cephalotho''ax is wide. There are two teeth on the 
inner margin of the falx-sheath, and three smaller on the outer. 

The stemium is broad shield-shaped, straight in front and 
curving to a point between the rear coxse, which are not quite con- 
tiguous. It is thickly covered with downlying fiat hairs and 
upstanding bristles. 

The Up is broader than long, rounded at the side and in front, 
where it has a broad sloping edge. 

The maxlllce are convex, upright, and about three times the 
height of the lip. Fiom their base they curve outwards on both 
inner and outer margins to a rounded upper end, so that they 
are broadest in the middle. They have a few short upright 
bristly hairs. 

The legs are moderately fine, covered with long powerful spines. 
There is a thick scopula on tlie tarsus a,nd metatarsus, and claw- 
tufts under the weak curved pair of claws. On tibii^e 1 and 2 



308 MR. H. R. HOGG ON SOME 

are three pairs of spines on the under side, on metatarsus of same 
a pair of very long spines at tlie base underneath, and .another 
about the middle at the side. On tibia 4 are three long spines on 
the upper side and three pairs on the under. On metatarsus 4 
are two pairs on the under side, three long spines on the outer 
side, and three long single ones on the upper side. 

The tibia and patella of the palp are of equal length. 

The measixrements (in millimetres) are as follows : — 

Long. Broad. 

Cephalothorax ... 3| i -^^ 

Abdomen 5^ 4 

Mandibles ij 

Pat. & Metat. 
Coxa. Tr. & fem. tib. & tars. 

Legs 1. li 4 4i 4i = 141 

2. 1| 4 4^4^= 14i 

3. 1 3 3| 4~ = 111 

4. U 5 5 Qi = 18 
Palpi I 2 2 l| = 6 

One female from Mt. Peel, Canterbury, collected by Mr. R. N. 
Hawkes. 

This species would appear to be somewhat near M. Simon's 
A. vittatus from Noumea, but the median abdominal stripe 
extends the whole length, and is pale brown instead of intense 
black. The legs are more heavily bespined, and the femora, 
although streaked longitudinally, are not ringed with black. 

Mynoglenes chiltoni, sp. n. (Text-fig. 95.) 

Male. The cephalothorax is of a i-ather deep yellow-brown, 
darker brown in the striations, but paler again on the rear slope. 
The eyes are pale yellow. The mandibles bright yellow-brown 
with red fangs. Lip and maxillse about the same as the man- 
dibles. The sternum darker, and dingier, is covered with fine 
granulations. The legs are pale yellov/ all over, with fine brown 
hairs paler towards the extremities. 

The abdomen, both on the upper and lower sides, is black-brown, 
scantily covered with fine reciimbent brown hairs. 

The cephalic part of the cephalothorax is slightly raised above 
the thoracic and rounded in front, where it is one-half the width 
of the broadest part, the latter being two-thirds of its total length. 
The surface of the thoracic part is smooth, without hairs, very 
finely granulated. The central fovea is shallow and round, with 
four pairs of small latei'al furrows, and a broader and deeper pair 
round the margin of the cephalic part, all radiating from the 
fovea. 

The eyes of the rear row are equal, the median their diameter 
apart, and the laterals one and a half diameters therefrom ; these, 
with the front laterals touching them, lie on prominent tubercles. 



NEW ZEALAND SPIDERS. 



309 



The front median eyes, half the diameter of the others, are one- 
half of their own diameter apart; they are a,s far from the rear 
median as the latter from the side. The chjpeus is as broad as 
the median eye-area is long, and the deep indentation characteristic 
of this genus runs below the front row of eyes. 

The mandibles are much swollen in the upper half, whence 
they narrow and become almost cylindrical in their lower half ; 
they are as long as the front of the head part is broad. On the 
outer margin of the falx-sheath are two long teeth followed by 
two smaller. 

Text-fip-. 95. 




Mynoglenes chiltoni. 

a, female ; b, male ; c, ej'es ; d, male palp from the front ; 
tlie side ; /, epigyue ; ff, lip aud maxillte. 



male palp from 



The li]) is broader than long, round at tlie top, with the pro- 
truding upper margin of the genus, and less than half the height 
of the maxilla2. These are ujoright, small at the base and gradually 
widen out as they curve over the lip. 

The sternum is a broad shield-shape, straight in front, rounded 
at the sides and running to a point at the rear, where the coxfe 
do not quite meet. The latter are rather unusually large and 
rounded, especially the anterior pair. 

Proc. ZooL. Soc— 1911, No. XXI. 21 



310 



ME. H. R. HOGG ON SOME 



The trochantal joints of the legs are long, and have a pronounced 
protruding ridge round them, about the middle. The other joints 
are thin and tapering, the tarsus and metatarsus being quite 
fine ; they are well covered with very fine greyish-brown hairs, 
but have no spines. 

The ahdomen is long and oval, the surface being much corru- 
gated longitudinally. The spinnerets are short, with a small 
hemispherical second joint. 

Except in size the female agrees with the male. 

The measurements (in millimetres) are as follows : — 

Larger Male. 
Long. Broad. 

Cephalothorax ... 3 -^ ^ 

Abdomen 4 2 

Mandibles 1 1 

Pat. & Metat. 
Coxa. Tr. & fem. tib. & tars. 

Legs 1. I 3 3 3i = 10,^ 

9 3 9-3 93 q _ Ql 

3. I 2| 2 2| = ^ 

4. I 3 3 4= 10| 
Palpi I n I ■ I = 3i 

Female. 
Long. Broad. 

n 1 1 4.1, o f 1 in front. 

Cephalothorax ... A \ '[i 

Abdomen 2 1| 

Mandibles 1 

Pat. & Metat, 
Coxa. Tr, & fem. tib. & tars. 

Legs 1. I n 2 1| = 6| 

2. ^ 2 H U = ^ 

.5. 2 ■'2 -"-2 ^2 — '^ 

4. ^ 1| 2 1=5 

Palpi i 11 1=3 

This species differs from M. marrineri in the paler colouring of 
the mandibles, lip, maxilhe, and legs. There is no yellow pattern 
on the back, only a slight mottling, and no paler area on the under- 
side of the abdomen as in the foregoing. The thoracic fovea is 
short and shallow instead of being long and deep, and the 
granulated portions are confined to the thorax and sternum. 

One male from the summit of Mt. Ngauruhoe, Waikato, N. 
Island. Mr. E. Phillips Turner. 

One male from Ruakura. Miss B. D. Cross. 

One female from Picton in the South Island, found by Prof. 
Chilton, after whom I have named the species. 



NEW ZEALAND SPIDERS, 



311 



DOLOMEDES TRIDENTATUS, sp. n. (Text-fig. 96.) 

Female. The cephalothorax is a bright chestnut, a broad area 
behind the eyes being thickly covered with silvery grey hairs, 
from this a narrower median streak of similar hair reaches to the 
rear slope, on each side of this is a broad area of chestnut 
covered with short orange hairs, and thence nearly to the side 

Text-fiff. 96. 





Dolomedes tridentatus. Female. 
a, eyes ; h, profile ; c, mandible fi-om inside ; d, tarsal claws ; e, epigjnie. 



margin is again a broad white-haired streak. The mandibles are 
dark red-brown, thickly covered with long bristly hairs, nearly 
white on the outer sides, darker grey on the inner. The fangs 
are red. The lip, maxillae, and sternum are dark brown, thickly- 
covered with long upstanding brown hairs. The legs and palpi 
are bright orange, with long brown hairs darkening towards 

21* 



312 MR. II. E. nOGG ox SOME 

the extremities of the limbs. The muscle parts are bright white. 
The abdomen is bright yellow-brown, thickly covered with short 
dingy yellow-brown hairs, a paler median stripe reaching from the 
base to about halfway down the back ; on the sides of the abdomen 
the hairs are white. The underside is quite dark yellow-brown, 
with four longitudinal white-haired rather narrow stripes. The 
spinnerets are bright yellow-brown. 

The upper sui'face of the cephcdothorax although flat is con- 
siderably raised above the level of the side margins, and slopes 
steeply at the rear. It is one-fourth longer tha,n its breadth in 
the widest part, and the front is only half the width of the 
latter. It is rounded at the sides and rear. There is a long and 
deep median longitudinal fovea extending from the cephalic part 
to the rear slope. 

The side eyes of the front row are on the lower slope of black 
tubercles, which, extending farther back than the median ej'es, give 
the appearance from above of a recurved row. Their lower edges 
are on a level with the lower edges of the median, and as they are 
smaller the line of the upper margins is really procurved. The 
median eyes are about their diameter apart and one-half that 
distance from the side pair, wiiich are two-thirds of the width of 
the former. The eyes of the second row are about twice the 
diameter of the median eyes of the fiont row. They are rather 
less than their diameter apart, the same distance from the front 
median, and the total length is clearly less than that of the front 
row. The median eye-area is broader than long. The eyes of 
the third row axe four-fifths the diameter of those of the second, 
five of their diameters apart, and two of the same from the eyes 
of the second row. The clypeus is about 2| times as broad as the 
front median eyes. 

The inandihles are kneed at the base, stout, as long as the 
front of Tihe cephalothorax is broad, and slightly divergent. They 
are thickly covered with long upstanding bristles. The fangs 
are broad at the base, strong, and Avell curved. About the 
middle of the inner margin of the falx-sheath are three lai^ge 
teeth, all equidistant, at the lower end of the outer margin 
is one large tooth between two small. The lip is broader than 
long, straight in front, and less than half the length of the 
maxillae. 

The sternum is a broad shield-shape, straight in front, pointed 
at rear, where it passes between the two rear cox«, which aie not 
quite contiguous. 

The legs are stout and straight, slightly and normally tapering 
to the anterior ends, but in nowise fine or flexible ; they are 
armed with numerous long and strong spines. There are scopulee 
on the tarsi and metatarsi of the front two pairs, on the tarsi 
only of the rear. The superior claws have six or seven teeth, the 
inferior are smooth. There is a pair of spines on patelJje of pairs 
3 and 4, none on patellie of 1 and 2. The femoral joint of the palp 



NEW ZEALAND SPIDEUS. 313 

is iiiciirved and enlarged at the anterior end, the tibial joint is 
longer than the patellar, and the female palp-claw has four pecti- 
nations. 

The abdomen is oval, rounded at the sides. The hairs are 
smooth and fine. On the upper side are three pairs of muscle- 
spots. The epigyne is horseshoe-shaped, hollow in the anterior 
half onlj, and without any longitudinal rib. The superior pair 
of spinnerets is longer than the infei-ior. They are both two- 
jointed, the second joint half the length of the first. 

The egg-sac is globular, white, and flocculent, 12 millimetres 
in diameter. 

The measurements (in millimetres) are as follows : — 







Long. 




Broad. 






Oephalothorax .. 


8 


{ 


f 3 iu front. 
1 6 




Abdomen 




11 

3 

Coxa. Ti- 


61 

Pat. & 
. & fern, tib 


Metat. 
& tars. 




Mandibles 












Legs 

Palpi 


1. 

2. 
3! 
4. 


3 
3 

^z 
3 


7 

7 

6i 

7' 

3 


7 
7 

8 
3 


H = 

6~ 

8 

2i = 


= 23| 
= 231 
= 21i 
26 
10 



One female from Dunedin. Collected by Mr, G. M. Thomson. 

This species differs from the accepted limits of the genus 
Dolomedes as detailed by M. Simon in having three teeth only 
on the inner margin of the falx-sheath instead of four. The 
clypeus while broad is less broad than the area of the median eyes, 
and the patellee of legs 1 and 2 are without spines. It is, however, 
much nearer to Dolomedes Latreille than to Thaumasia Perty, 
Drances Simon, or any other genus. I would have felt it right 
to propose a new genus for it on the above difi'erences but for the 
fact that it graduates so imperceptibly into other members having 
four teeth, notably D. huttoni and D. trippi from the Chatham 
Islands, which are certainly Dolomedes. Its eyes and clypeus, the 
equality of the 1st and 2nd pairs of legs, and the marking of 
the cephalothorax in the female correspond almost exactly with 
those of D. trippi. Its epigyne is of the same type as that of 
D. huttoni. 



314 MR. A. H. COCKS OX A 

EXHIBITIONS AND NOTICES. 

Fel)riiary 21, 1911. 

Dr. A. Smith Woodward, F.R.S., Yice- President, 
in the Chair. 

The Secretary read the following report on the additions made 
to the Society's Menagerie during the month of January last : — 

The registered additions to the Society's Menagerie during the 
month of Ja,nuary were 121 in number. Of these 63 were 
acquired by presentation, 17 by purchase, 32 were received on 
deposit, 5 in exchange, and 4 were born in the Gardens. 

The total number of departures dui-ing the month, by death 
and removals, was 160. 

Amongst the additions special attention may be called to the 
following : — 

2 Lion Cubs (Felis leo), from the Sebakwe River, S. Rhodesia, 
presented to the King's South-African Collection, through 
H.R.H. The Duke of Connaught, K.G., by Col. Weston Jarvis. 
Deposited by H.M. The King on Jan. 14th. 

1 Jaguar (Felis onca), born in the Gardens on Jan 24th. 

2 Bay Duikers (Cephcdop/ms dorscdis), from Coomassie, pre- 
sented by Capt. S. H. Chapin on Jan 21st. 

2 "Virginian Deer {Boi'celaphics americanus) $ $ , from ISToi'th 
America, purchased on Jan 17th. 

1 Kacunda Nightjar {Podager nactonda), captured at sea ofi' 
the coast of Brazil, new to the Collection, purchased on Jan 2nd. 

1 Bornean Fireback Pheasant [Lophura ignita), presented by 
H.G. The Duke of Bedford, K.G., Pres.Z.S., on Jan 21st. 



■ Dr. H. Hammond Smith, M.R.C.S., F.Z.S., exhibited three 
skins of male Pheasants assuming female plumage, sent to him 
by Mr. Arthur Gilbey, and some microscopical specimens of the 
glands prepared by Dr. S. G. Shattock. 



Mr. G. A. Boulenger, F.R.S., Y.P.Z.S., contributed a paper 
based on a collection of Fishes from the Lake Ngami Basin, 
Bechuanaland, made by Mr. R. B. Woosnam, F.Z.S. 

This paper has been published in the ' Transactions.' 



Mr. Alfred H. Cocks, M.A., F.Z.S. , exhibited a series of 
photographs of the female Brindled Gnu recently born in the 
Society's Gardens, and gave the following account of its growth 
and coloration :— 

" On receipt of the orphan calf of the Brindled Gnu, born in 



YOUNG BRINDLED GNU. 



315, 



the Gardens on the 1st December last, which was brought to me 
on the 3rd, I was struck by the great difference between her and 
the drawing by J. Smit in P. Z. S. 1900, Plate xlviii. (facing 
p. 771), representing another calf previously born here; that 
figure, and one in ' Illustrations of the Zoology of South Africa,' 
&c., by Andrew Smith, M.D., London, 1849, No. 16, Aug. 1842, 
Plate xxxvii. *, being, I believe, the only two existing of a Gnu 
calf of this species. 

Text-fio-. 97. 






'^^'^^""''f 



-'^ 





Young Brindled Gnu bom in the Societj^'s Gardens on December 1, 1910. 
From a pbotograpb taken on January 7, 1911. 

" What seemed sjiecially worthy of note was that the tail for 
the proximal two-thirds of its length was white, as if the white- 



* The beginning of A. Smith's description of the young is quaint: "Form and 
appeai'ance clumsy and unseemly " ! 



316 



MR. A. H. COCKS ON A 



tailed species were the older form. The body-colour was also 
quite different from that shown in Smit's plate, being of the lustre- 
less ash-brown of an immature house-mouse, with a dull or rusty 
black dorsal stripe terminating in a point at the sacrum. The 
cheeks instead of being smooth and ma.tching the body in colour, 
as shown in both the above-mentioned plates, were exactly the 
reverse. The dark colour on the outer side of the ears did not 

Text-fig. 98. 



^ 



"----, 




\ 



.>^' 



Head of young Brindled Gnu. 
From a pliotograph taken on Januarj' 7, 1911. 



quite reach the margins, as if a man had taken a brushful of 
paint, and had given one streak to each ear, without afterwards 
making good the deficiencies. The lower two-thirds of all four 
legs was quite white ; Andrew Smith's plate shows this con- 
spicuously. The whole face was very dark or black, the black 
extending to a sharply defined width of about half an inch round 
the lower side of the eyes. 



I 



YOUNG BRINDLBl) GNU. 317 

" By the 13th December (that is, when the calf was twelve days 
old), the whole of the upper side of the body was a light fawn (the 
original 'puppy coat' having been shed) ; the legs were gradually 
colouring, or less pure, or conspicuously white, than at first ; and 
the colour was extending from above downwai'ds. The white on 
the tail was already nearly gone. Face dark ; nose and muzzle, 
as before, black ; cheeks quite light, the dark ring remaining 
round the eyes. The coloration of the exterior of the ears as 
before. 

" In Smit's plate the fawn of the body is darker, and more 
mixed with black-tipped hairs, and the drawing was evidently 
made at a greater age* than the present example attained i;o, 
not only for the reason just stated, but because the tail-tuft is of 
a brighter black, the dark circle round the eyes is shown as the 
faintest possible shadow, and the general figure of the animal 
is more set ; but the most important discrepancy — which seems 
only explicable on the supposition that the present example be- 
longed to a local race widely difiering from the two calves previously 
figured, — is that both plates show quite a smooth cheek, whereas 
the present example, like her father (and I feel sure the mother 
was the same), had a perfect and veiy conspicuous ' Newgate 
fringe,' consisting of a woolly moustache, like a poodle's, but 
situated about halfway along the gape, and quite distinct from 
the moustachial bristles on the muzzle, meeting bushy whiskers 
on the cheeks, and continued as a long thick beard on the chin ; 
the whole cheek was hirsute, or covered with long hairs projecting 
in a more or less upward direction. 

" By the 16th the horns were appearing through the skin, and 
she had cut some molar teeth, which we could hear her grinding, 
so we tied up a wisp of hay, and she very soon learnt that it was 
good to eat. The tail now much resembled that of the Grey 
Squirrels so familiar in the Pai-k, with the addition of a black tuft. 

"By the 27th the dark ring below the eyes was fading 
away. 

"On the 14th Januai-y the horns were about an inch long ; and 
on the lower surface of the chin, between lip and Newgate 
fringe, but hardly extending to the side face, was a white patch, 
which I had not previously noticed. 

"On the 15th the calf appeared as vigorous and in as perfect 
health as ever, bxat on the morning of the 16th she lay prostrate, 
and in a very short time was dead ; the cause, as decided by 
Mr. Plimmer, the Society's pathologist, being broncho-pneumonia. 

"The Gn\i wns photographed, by myself, on December 13, on 
December 31 and January 2, by Press photographers, and on 
January 7, by request of Dr. Chalmers Mitchell, by a Henley 
photogi'apher ; the series to some extent shows the growth. 

* No hint is given (loe. cit.) as to the age of the calf when drawn, but it was born 
en July 14, and the drawing exhibited on November 20. 



318 MR. H. G. PLIMMER ON DEATHS 

" I took tlie following measuremeBts immediately after death : — • 
Extreme length, tip of nose to tip of tail ( = vertebrse), 4ft. 1| ins. 
Length of face, 11-1- ins. 

,, ear, 5| ins. 

., tail (to end of vertebrae), 11 ins. 

,, fore leg (elbow to end of toe), 22g ins. 

,, hind leg (hock to end of toe), 15^ ins." 



PAPERS. 

15. Report on the Deaths which occurred in the Zoological 
Gardens during 1910. By H. G. Plimmer, F.R.S., 
F.Z.S., Pres.R.M.S., Pathologist to the Society. 

[Received February 6, 1911 : Read February 21, 1911.] 

On January 1, 1910, the number of animals in the Zoological 
Gardens was 3186, and during the year 2354 animals were 
admitted, making a total of 5540 for the year. 

The number of deaths during the year has been 1554, that is 
a death-rate of 28 per cent. ; but if from the above total we 
deduct 643 animals which did not live for six months after their 
arrival in the Gardens — that is, roughly, the time at which we 
find they have either got over their journeys, or died from any 
disease they brought with them, or have got entirely used to their 
new environment — the percentage of deaths is reduced to 16'4, 
which is less than that of the last two years. 

The following figures will show the general relations of the last 
three years : — 

1908. 1909. 1910. 

Total number of animals 5608 5303 5540 

Total deaths 1737 1492 1554 

Percentage of deaths 31 28 28 

Percentage of deaths, excluding 
those which occurred in ani- 
mals which had oiot been six 
months in the Gardens 17 17-8 16-4 

If we consider the weather conditions of 1910, and compare 
these figures (bearing in mind the relative number of animals), 
it will be seen that the percentage of deaths of 1910 is more satis- 
factory than that of the two preceding years. 

The total deaths are divided as follows : Mammals 362, Birds 
845, Reptiles 347. 

The following tables show the facts ascertained in outline. 
Table I. sets forth the actual causes of death in each of the three 
groups specified. Under Reptiles are included Batrachians and 
Fishes. 



IN THE GARDENS DURING 1910. 

Table I. — Analysis of the Causes of Death. 



319 



Disease. 



Microbic or Parasitic 
Diseases. 

Tuberculosis 

Mycosis 

Pneumonia 

Septicsemia 

Abscess 

Pleuritis 

Empyema 

Pericarditis 

Peritonitis 

Stomatitis 

Hydatids 

Worms 

Hsemogregarines 

Malaria 

Coccidiosis 

Leucocytozoon infection . 

Cj'stitis 

Pancreatitis 



Diseases of Respiratory 
Organs. 

Bronchitis 

Broncho-pneumonia 

Congestion of lungs 

Atelectasis 



Diseases of the Heart. 
Dilatation of heai-t . . . 
Fatty degeneration ... 

Diseases of Liver. 
Patty degeneration ... 
Hepatitis 



Diseases of Alimentary Tract. 

Over-distension of crop 

Gastritis 

Gastric ulceration 

Gastro-enteritis 

Enteritis.. 

Intussusception 

Intestinal obstruction 



Diseases of the TJri^iary and 
Reproductive Organs. 

Nephritis 

Stone 

Inflammation of oviduct 

Diseases of Nervous System. 
Myelitis '. 



Mammals. 



Yarious. 

Carcinoma 

Sai'coma 

Puerperal eclampsia 

Senile decay 

Rickets 

Leutamia 

An.-smia without ascertained 7 

cause 5 

Injuries discovered post- \ 

mortem ( 



18 

13 

31 

7 

1 

1 



3 

50 
41 

2 



19 

32 

6 



22 
1 



Birds. 


Reptiles. 


163 


9 


99 


6 


73 


82 










6 


"i 


2 




2 


3 


"9 


"6 




16 


8 






1 


1 




i 




112 


14 


"7 




15 


1 


4 


1 


2 






1 


i 




3 


10 


186 


27 


2 




12 




'3 




1 




2 




"1 




1 




1 




1 




16 


2 



Reference 
to Notes 
following. 



10 
11 



12 
13 
14 



15 



16 
17 



18 



320 



MR. H. G. PLIMMER ON DEATHS 



Besides those tabulated above, 

113 animals were killed by order or by companions, 
101 ,, died of starvation and malnutrition, 

7 ,, were not exainined, 
108 ,, were too stale for detailed examination, 

these completing the total. 

In Table I. the classification is made into those diseases which 
actually caused the death of the animals, but in a large number 
of Mammals, Birds, and Reptiles other lesions were present which 
helped towards the fatal issue, and these are classified in Table II. 
If taken together with Table I. a much more accurate estimate of 
the amount of disease in the Gardens will be arrived at. 



Table II. — Other Diseases found in the Animals tabulated in 

Table I. 



Diseases. 


Mammals. 


Birds. 


Reptiles. 


Reference 
to Notes. 


Tuberculosis 


12 
1 
6 

2 

8 

8 

14 

1 

"2 
4 
1 

2 
5 

26 

12 

3 

4 

22 
1 
5 
1 

"6 

12 

13 

i 

23 


34 

8 

15 

"2 
3 

2 

12 

7 

36 

51 

48 

6 
3 

48 
3 
1 

1 

29 

3 

14 

7 

"2 


8 

5 

14 

"3 

1 

21 

'5 

"4 

10 
1 

i 

2 

"3 

"5 
13 


19 
20 


Mycosis 


Pneumonia 


Stomatitis 


Worms 


Peritonitis 


Pericarditis 


Empyema 


Trypanosomes 


Hsemogregarines 


Malaria 




Pleuritis 


Filariifi 


Cystitis 




Bronchitis 




Congestion of lungs 


Collapse of lungs 


CEdema of lungs 


Hvdrothorax 


Dilated heart 


Fatty heart 


Fatty liver 


Hepatitis 


Gastritis 


Gastric ulceration 


Gastro-enteritis 


Enteritis 


Intestinal obstruction 


Nephritis .. , 




Osteo-arthritis 


Rickets 





IN THE GARDENS DURING 1910. 



321 



Table III. — Shows the Distribution of Diseases causing Death 
amongst the principal Orders of Mammals. 



Diseases. 



Tuberculosis . 
Mycosis .... 
Pneumonia . 
Septiciemia . 
Pleuritis .... 

Abscess 

Empyema . . . . 
Pericarditis . 
Peritonitis . 
Hydatids .... 

Worms 

Cystitis 



Bronchitis 

Broncho-pneumonia 
Cong'estiou of lungs 

Atelectasis 

Dilatation of heart . . 

Fatty heart 

Fatty liver 

Hepatitis 

Gastritis 

Gastric ulceration .. 

Gastro-enteritis 

Enteritis 

Intussusception 

Nephritis 

Stone 

Myelitis 

Carcinoma 

Sarcoma 

Eclampsia 

Senile decay 

Rickets 

Injuries 



15 
16 



4 

1 

12 

3 

7 



3 

15 



10 



10 



6 

4 



As Tuberculosis and Mycosis have again been the cause of a 
large number of deaths amongst the Birds, the following Table 
has been drawn up to show the relative incidence in the various 
Orders. 

Table TV. — Showing the Com.parative Incidence of Tuberculosis 
and Mycosis in the various Orders of Birds. 



Orders. 


Tuberculosis. 


Mycosis. 


Passeres 


69 

30 

12 

2 

16 

19 

12 

1 

2 


22 

17 

6 

8 

6 

16 

18 

5 

1 








Columbee . 


Gallinae ... 


Striges 


Laridse 


Struthiones 





322 ON DEATHS IN THE GARDENS DURTNG 1910. 

Notes on the foregoing Tables. 

The following Notes refer to some points of special interest 
which it was not possible to include in the tables : — 

1. It will be noticed that there has been about the same 
percentage of tubercle as last year amongst the Mammals and 
Reptiles ; that is, it has been found in 10 per cent, of the 
Mammals and 4"9 per cent, of the Reptiles examined. In the 
Birds, however, there has been an increase of 107 cases; that is, 
it has been found in 24'9 per cent, of those examined. This 
increase amongst the Birds has been partly due to an epidemic 
which occurred in the ISTew Bird House in the beginning of the 
year. This was dealt with by entire disinfection of the house, 
and some structural alterations Avhich have greatly improved it ; 
a very definite improvement has taken place, but the arrange- 
ment of the cages at the ends of the house, and overcrowding 
generally amongst the Birds, is a constant source of danger. 

Amongst the Mammals 2 cases have been of bovine type, and 
amongst the Birds there have been 7 cases of this type. 

The liver of a Tantalus showing this particular type has been 
sent to the Museum of the Royal College of Surgeons. 

2. Mycosis (under which term several distinct mould-diseases 
are grouped) has also increased among the Birds to a considerable 
degree — 107 cases as against 48 last year. 

The Mammals dying from this disease have been principally 
Wallabies and Gazelles. The disease (formerly grouped under the 
Septicaemias) has been found to be caused by a mould which enters 
through the mouth and produces abscesses about the jaws and in 
the adjacent muscles, and eventually a septicaemia. 

It has also been found that a disease of the eye in Birds is due 
to a mould. It commences as a keratitis, and then pus is 
formecl in the ai^terior chamber, and secondary abscesses form, 
from which the bird dies. Of the division headed Reptiles 
3 were fishes with a mould -disease of the skin which invaded the 
muscles. 

3. The increase of pneumonia amongst the Reptiles (87 cases as 
against 64 last year) has been mainly due to the variable 
temperatures in the Reptile House, owing to alterations in the 
heating apparatus. Only 3 of these cases were due to the presence 
of worm-eggs and embryos. There has been a decrease amongst 
the mammals and birds. 

4. Five of these cases followed injuries : the remaining two 
were in Wild Swine, where it was caused by an organism beloiig- 

' ing to the group producing HaBmorrhagic Septicaemia. 

5. The one Mammal was a Fox with suppuration in the middle 
ear and destruction of the semicircular canals. 

6. In an Antelope the pericarditis was due to numbers of 
worm-cysts on the visceral pericardium, which, so far as I know, 
is unique. In the Birds it was mostly due to the deposition of 
uric-acid crystals in the pericardium. 



I — I 
I — I 
en 

m 

N 




ox THE VARIATIONS OF THE SPOTTED SALAMANDER, 323 

7. In the Mammal — an Aardwolf — the worms were encysted 
pentastomata, which were in large numbers all over the body, 
causing pleuritis and peritonitis. In the Birds they were mostly 
syngamus. 

8. Under Malaria are grouped cases in which intracorpuscular 
parasites belonging either to the Halter klkmi or Proteosoma 
group have been found in sufficient numbers to cause death. 

9. In a Bulbul in which 60 per cent, of the polynuclear leuco- 
cytes were infected. This I believe to be new. 

10. Most of these occurred during the first and last three 
months of the year ; 4 of them were badly rickety. 

11. Nine of the Mammals had bad rickets also. The number 
of cases amongst Birds, in which it is relatively much more fatal, 
is slightly less than last year. 

12. In most of the Mammals it was caused by food-jjoisoning, 
in others, and in the Birds and Reptiles, by worms burrowing into 
the mucosa. 

13. In 7 Mammals, 59 Birds, and 1 Reptile the enteritis was 
hsemorrhagic ; in 6 Birds and 6 Reptiles it was due to woi-ms ; and 
in 9 Birds and 1 Mammal it was due to foreign bodies. This 
disease is a little less prevalent than last year. 

14. Two very extreme intussusceptions occurred in two Wom- 
bats, recent arrivals, which came together. 

15. In a Coypu Rat, in which 27 stones were found. 

16. Four of these cases of cancer were in Wallabies, the 
stomach in all was the seat of the primary growth ; the fifth was 
in a Markhoor in the mouth. 

17. Two Gazelles died from sarcoma, one of liver and one of 
mediastinal glands. 

18. This occurred in a Jungle-fowl and was of the spleno- 
medullary variety. 

19. In a Partridge and Mai^sh-Bird, both not described before. 

20. In a Bat, in which all the wing-joints Avere affected. 

16. A Contribution to the Study of the Variations of the 
Spotted Salamander (Salaynandra tnaculosa). By 
Edwaed G. Boulenger*. 

[Received December 10, 1910 : Read February 21, 1911.] 

(Plate XY.t & Text-figures 99-102.) 

The experiments now being carried out in Vienna by 
Dr. Kammerer on the colour-changes of the Spotted Salamander 
[Scdamandra maculosa) in relation to its environment are 
attracting attention, and it has occurred to me that a general 
survey of what is known of the varieties of this very variable 
species, especially in connection with the geographical distribution, 

* Communicated \iy G. A. Boulengeb, F.R.S., V.P.Z.S. 

•f- For explanation of the Plate see p. 3t7. 



324 MR. E. G. BOULE>fGER ON THE 

would not be without importance at the present moment. Such 
a survey would afibrd those who wish to follow the path opened 
lip by Dr. Kammerer a more precise basis than can be obtained 
from the available literature on the subject. I have therefore, 
with the help of my father, undertaken a revision of the rich 
material in the British Museum and have now the honour of 
offering an account of it for piiblication to the Zoological Society. 
One of the jDrincipal results of my study has been to lay greater 
stress on the disposition of the markings than on their actual 
size, form, or colour, and to define two principal forms in Central 
Europe, which previous authors have not separated with sufficient 
precision, in spite of their well-marked geographical segregation. 

Except in the case of var. molleri, with its aberrant coloration, 
authors dividing S. maculosa into a number of subordinate forms 
have dwelt on real or supposed structural characters, whilst 
ignoring the disposition of the markings. Bedriaga (3, p. 98), 
the most recent writer on the subject, recognizes, apart fi'om the 
typical form, also called by him var. europcea (2, p. 252), three 
varieties, namely — var. algira, var. Corsica, and var. molleri, the 
two former based only on slight structural differences, which are, 
besides, not constant, as I intend to show further on. Under the 
designation of typical form authors have generally thrown together 
specimens with different styles of markings, either simply ob- 
serving that these are subject to infinite variation or classifying 
them under a number of titles, which refer merely to indi- 
vidual variations, such as the var. tceniata, var. quadrivirgata and 
var. nigriveniris, j^roposed by Diirigen (10, pp. 577 & 578) for 
certain individuals. In order to avoid introducing a new name, 
I will adopt the first of these for the assemblage which I have 
endeavoured to define and contrast with the typical spotted form 
on which the name maculosa is based. 

To better bring out the individual differences to which the 
markings are subjected in this Salamander, I have drawn up 
ta.bles of a certain number of the specimens in the British 
Museum, upon a scheme which should prove of use to those 
making experiments on the colour-changes, as by that means a 
record of each individual specimen, out of a large nmnber, can be 
kept in such a way as to ensure its future identification. Such 
tables, explained by the annexed diagram (text-fig. 99), do not, 
however, convey an exact representation of the markings, which 
can only be done by descriptions, but they will be found to answer 
well enough for the purpose of identification. 

In defining the varieties into which the species Salamandra 
macidosa may be divided, I have not lost sight of occasional 
exceptions, and have duly pointed them out. There are always 
exceptions, especially when we have to deal with forms of sub- 
specific rank, but such as I have come across are too few to 
militate against the adoption of a var. iceniata as opposed to the 
forma typica. 

In the tables the explanation of the various columns is as 



VARIATIONS OF THE SPOTTED SALAMANDER. 



325, 



Text-fiff. 99. 



follows : — The length of each specimen (in millimetres) is taken 
from the end of the snout to the posterior extremity of the vent. 
O means presence ( + ) or absence ( — ) of the supraocular spot 
(on the upper eyelid). 

P, the spot on the parotoid gland. 

S, spot or spots on the snout. 
OP, OS, whether the above- 
mentioned spots are confluent or 
not. PB, whether or not the 
spot on P is confluent with one 
on the body. In cases where the 
two sides differ, the initials R 
(right) and L (left) indicate the 
discrepancy. 

Sp., total number of spots on 
the upper surface of body (to 
a line connecting the posterior 
borders of the hind limbs), which 
is divided into four conventional 
segments, numbered I, II, III, 
lY ; under each of the latter 
figures, the spots pertaining to 
each segment are given. These 
spots are numbered 1, 2, 3, &c., in 
the order shown on the diagram ; 
in case of absolute symmetry, the 
spots are numbered from right to 
left. 

A stands for the vertebral area, 

, _ J bearing the two median rows of 

\ ^ ^ ___ _' / glands. B (right and left) for the 

- ^ ' " area between A and ; (right 

and left) for the area occupied by 
the series of large lateral glands ; 
D for the area between the latter 
and the limit of the belly (taken 
from an imaginary line connect- 
ing the lower border of the axil 
with the groin). In column D, 
right and left sides are separated 
by a — , the figures referring to 
the number of spots on each side 
(spots confluent with the ventrals 
and dorsals not reckoned), not to 
the individual spots as in the 
columns A to C. 
G, pi-esence ( + ) or absence ( — ) of yellow spots on the chin and 
gular I'egion. 

Y, on the ventral region of the body. 




Diagram to explain the tables of 
different markings in Salamandra 
maculosa. 



Proc. Zool. Soc— 1911, No. XXII. 



22 



326 



MR. E. G. BOULENGER ON^ THE 



The tabulation of tlie markings on the specimen represented in 
text-iig. 99 may be drawn up as follows : — 



Section. 

O OP P PB S OS Sp. iT II. III. IV. C B 
+ _+___ 6 1 1.2.3 4.5 6 - 1.2.4.6 



Ake^i 



A B C 

1.2.4.6 1.3.4.5.6 3 



The following is a list of the specimens preserved in the 
British Museum and the Lataste Collection, arranged geographi- 
cally under varieties : — 



Forma typica. 



France. 



1. 

2. 
3. 

4. 




Renre, near Besan^on. 

Avignon. 

Basses-Alpes. 

S.E. France. 

Corsica. 


Mme. Phisalix. 

M. M. Mourgue. 

M. Honnorat. Lataste 

Collection. 
Prof. Buboscq. 


5-9. 

. 10. 

11. 


? & yg- 

Yg. 
Yg. 


Bocognano. 
Vizzavona (1100 m.). 
Corsica. 

Germany. 


Dr. J. de Bedriagn. 
Prof. Vayssifere. 
Di-. J. de Bedriaga. Lataste 
Collection. 


12. 
. 13. 
14-15. 




Near Stuttgart. 
Ballenstadt, E. Harz. 
Sharfeiiberg, near Meissen, 


Prof. K. Lampert. 
Dr. W. Wolterstorff. 



16-17. <??. 

18. Hgr. 

19. S. 
■ 20. $. 



21. c?. 

22. ?. 
23-24. ?. 

25-54. <?. 

55. <?. 

56-60. S ? , 

61. ?. 



62-63. $ ? . 

64-66. d' ? & ys 

67. ?. 

68. Yg. 
69-88. <??. 

89. <?. 

90. ^. 
91-92. <?. 



Saxony. 



Switzerland. 



Near Lausanne. 
Fliielen, Uri (465 m.). 
Lugano, Ticino (275 m.). 
Val Bavano at Cavergno, 
Ticino (ca. 600 m.). 

Italy. 

Prego, Brianza, L. Como. 
Tuduno Olona, Varese. 
Olginate, Prov. Lecco, 

Lombard}\ 
Cascinella, near Borgoli, Prov. 

Genoa. 
Viterbo. 
Prov. Rome. 
Aspromonte, near Reggio, 

Calabria (1600 m.). 

A ustria-Hungary. 

Hiitteldorf, near Vienna. 

Near Prague. 

Brasso, Hungary. 

Teszla, Bozau Mts., Hungary. 

Nagy Becskerek, Hungary. 

Hungary. 

Sarajen, Bosnia. 

Travnik, Bosnia. 



W. Morton, Esq. 
M. A. Gliidini. 



Dr. C. Vandoni. 



Dr. R. Gestro. 

Prof. J. J. Bianconi. 
Prof. Carruccio. 
Prof, 0. Neumann. 



Dr. F. Werner. 

G. A. Boulenger, Esq. 

Prof. L. V. M^hely. 

Hr. A. V. Kovdcs. 

Dr. F. Wernei-. 



VARIATIONS OF THE SPOTTED SALAMANDER. 



327 



Roumania. 

93. c?. Near Azu, Carpathians of M. A. MontHndon. 

Moldavia. 
94-113. (J 9 <^ 3'o- Siiiaia, Carpathians of Vallachia. „ 



Ilk Yff. 



115. ?. 



Parnassus 



Greece. 



Asia Minoi\ 



Dr. T. Kriiper. 



Zebel Bulgar Dagh, Cilician C. G. Danford, Esq. 
Taurus (1200 m.). 



Algeria. 

116-118. ? & yg. Mt. Edough, near Bona. 
119-122. <? $ & yg. 



123. Yg. 

124. ?. 
12-3. <?. 



L'Arba, near Algiers. 
Algeria. 



Morocco. 
126-133. ? & yg. Benider Hills, near Tangier. 



Dr. Hagenmiiller. 

Lataste Collection. 
M. L. Bedel. 
M. Lallemant. „ 



M. H. Vaueher. 



Vars. GALLAICA and molleri. 

Spain. 

1. $ . Vigo, Galicia. M. V. L. Seoane. 

2-5. cj ? & yg. Galicia. „ 

Lataste Collection. 
6-11. d' ? & yg. Loroya Valley, near Madrid M. de la Escalera. 
(300-1.00 m.). 
12. Yg. Spain. Lord Lilford. 



Portugal. 



13. ?. 

14-17. c??. 

18-19. S ? ■ 

20. ?. 



Coimbra. 



Ciutra. 
Near Lisbon. 



Dr. J. de Bedriaga. 

Col. Yerbur3\ 

Sr. Mattozo Santos. 



Var. T^NIATA. 



France. 



1. 


?. 


Near St. Malo. 


G. A. Boulenger, Esq. 


2-5. 


(??. 


lloscoft', Finistere. 


„ 


6-8. 


? & yg- 


Near Rouen. 


M. Louis Miiller. 


9. 


?. 


,, 


M. L. Horst. 


. 10. 


?• 


Armainvilliers, near Paris. 


M. E. Simon. Lataste 
Collection. 


11-13. 


<?. 


Haute-Marne. 


Dr. A. Pettit. 


14-19. 


<??• 


Beure, near Besan^on. 


Mme. Phisalix. 


20. 


?. 


Bourg-en -G ironde. 


Lataste Collection. 


21-26. 


<??• 


Herault. 


Prof. Duboscq. 


27. 


Yg. 


Aix-les-Thermes, Ariege. 


M. V. Baillet. 


28. 


$. 


Eaux-bonnes, Basses-Pyrenees. 

Belgium. 


Rev. F. A. Walker. 


29-30. 


<??• 


Maredsous, Prov. Naniur. 


Rev. B. Lebbe. 


31. 


(?. 


Waulsort, „ 


G. A. Boulenger, Esq. 
22* 



328 MR. E. G. BOULENGER OX THE 







Luxemburg. 




32. 


?. 


Luxemburg. 

Germany. 


M. V. Perrant. 


53-40. 


(? ? & yg- 


Ilsenberg, Harz. 


Dr. W. Woltcrstorff. 


41. 


?. 


J 


W. H. Decks. 


42-51. 


^?- 


Harz. 


Zoological Society. 


52-63. 


Yg. 


^j 


G. A. i5oulenger, Esq, 


54-55. 


(??• 


Yorwolile, Brunswick. 


Dr. W. WolterstortV. 


56-58. 


<??. 


Holzminden, Brunswick. 


„ 


59-60. 


(??• 


Stadtoldendorf, Brunswick. 


M. A. Ghidini. 


61-63. 


(? &yg- 


Lippe-Detmold. 


Dr. J. Roax. 


64-67. 


<? ? & yg- 


Near Stuttgart. 


Prof. K. Lampert. 


68. 


(?. 


G. Duchy of Baden. 

Switzerland. 


Basle Museum. 


69-72. 


<? 9 & yg- 


Langenbruck, Jura. 


Basle Musenm. 


73. 


^- 


St. Gallen. 

Portugal. 


M. A. Ghidini. 


74-75. 


^ & yg- 


Oporto. 


E. Allen, Esq. 


76-77. 


<?. 


Portugal. 


P. B. Webb, Esq. 



I. The Typical Form. 

This form deserves to be regarded as the typical, not only 
because the name maculosa applies best to it, but because it is the 
only one found in Austria {cf. Werner, 32, p. 119), where it was 
described under that name by Laurenti (18, pp. 42, 151). It also 
happens to be the form figured by most authors :— Aldrovandi(l, 
p. 641), Rosel (24, frontispiece), Latreille (17, pi- i-), Sturm (30), 
Reider & Hahn (23), Funk (13, pi. 1.), Bonaparte (5), Rusconi 
(25, pi. i.), and Camerano (6, pi. i.). It corresponds to the vars. 
A and C of Dumeril and Bibron (9, p. 37j, a-c of Schreiber (27, 
p. 75). In this form the black nearly always greatly predominates 
over the yellow, the latter appearing as markings of various 
shapes, — round, elongate, C". S-, Y-shaped, &c., and disposed over 
the body, often in 3 to 5 alternating series, or with a median 
series forming a sinuous or zigzag vertebral stripe. If, as is very 
exceptionally the case, the dorsal spots appear to form two longi- 
tudinal series, it will be observed that they by no means hang 
together in regular chains continuous with the spots on the 
parotoids. Only in one specimen (from Lausanne) have I felt 
embarrassed as to the form to which it should be referred. Upper 
eyelid and parotoid usually entirely, sometimes only partially, 
yellow, the spots on the parotoid may even be entirely absent 
(specimens from Algeria and Morocco). The two spots thus 
located are, as a rule, distinct, but may occasionally run together. 
In a specimen from ISTagy Begskerek, Hungary, the yellow 
markings on the upper eyelids extend across the interorbital 
region, forming a cross-bar. The sides usually bear spots, which 
may number vxp to 15, in which case they are, of course, very small. 
A spot above the angle of the mouth is absent in only about 5 per 



VARIATIONS OF THE SPOTTED SALAMANDER. 



329 



cent, of the specimens. Gular region with a few large or small 
markings, sometimes completely unspotted. Below entirely black 
or with spots which are usually of small size, the only specimens 
with very large spots being single ones from Genoa, Yiterbo, and 
Reggio in Italy. Snout generally entirely black. Limbs black, 
with a characteristic yellow blotch on the upper surface of the 
arm and thigh, near the base, which is constantly present ; a 
second on the forearm and leg may or may not also be present. 
Hand and foot entirely black or with one or two (rarely more) 
yellow spots, which, as a rule, are not confluent with the spot on 
the forearm and leg. Single or paired spots on the tail, which 
may run together to form a stripe. Under surface of tail nearly 
always black. 

Text-fig. 100. 





Forma tijpua. 



The coloui- of the bright markings varies from chrome-yellow to 
a deep orange. I have found traces of claret-red on the head in 
some specimens from Genoa. The usual absence of yellow on 
the snout has been mentioned above ; the only exceptions I have 
noted are to be found in specimens fi'om Lausanne, Avignon, 



330 TARIATIONS OF THE SPOTTED SALAMANDER. 

Corsica, Genoa (7 specimens out of 40), the province of Rome, and 
Calabria. The latter is remarkable for the large size of the yellow 
markings on the upper and lower surfaces, where they almost equal 
in extent the black area. The opposite extreme occurs in a spe- 
cimen from the Yal Bavano, Ticino, in which the yellow colour is 
reduced to a few very small blotches on the parotoids and at the 
base of the limbs. These two extremes are figured side by side in 
text-fig. 100. 

On careful examination of the specimens of var. algira Bedriaga 
and var. Corsica Savi, I have come to the conclusion that they are 
not essentially difl[erent from the forma typica. 

The var. algira is described by Bedriaga (3, p. Ill) as having 
the tail and digits longer and more slender than in the typical 
form ; but the specimens from Mt, Edough (in the Lataste 
Collection), upon which Bedriaga based his observations, are in a 
leather emaciated condition, having, no doubt, been kept in 
captivity for some considerable time, and this is evidently partly 
the cause of their slenderness. Although the digits and tails of 
the Salamanders from Algeria and Morocco are, as a rule, longer 
than in the typical form, the longest digits in the latter may be 
actually longer in proportion to the length of the body. Thus, 
in a specimen from Lake Oomo, the length of the longest toe is 
9 2 per cent, of the total length (from tip of snout to posterior 
end of vent), while in a specimen from Mt. Edough, Algeria, the 
length of the same is 9 per cent,, and in one from the Benider 
hills, Morocco, as low as 6| per cent. Again, the length of the 
tail in var. algira ranges from 65 to 81 per cent, of the length of 
the body, against 54 to 78 in the forma typica, an overlap which 
precludes the character being used as diagnostic. 

In the same author's description of the Corsican variety, the 
head is stated to be remarkably broad, and the toes to be much 
more strongly depressed and with sharper edges on the sides than 
in the typical form, I have examined the very specimen described 
by Bedrifiga, but do not find the head to be any broader than in 
some of the typical and striated forms, and although the toes are 
more depres^'ed than is generally the case, they are not more so 
than in certain specimens from Vienna, Bosnia, Luxemburg, and 
the Harz Mountains. As to the more sharply edged sides of the 
toes, this sharpness simply coincides with the degree of depression. 

The supposed difference in the palatine dentition, on which 
S. Corsica was founded by Savi (26), has long ago been disposed of 
by Schreiber (27), Bedriaga (2), and Camerano (6). 

The habitat of the typical form seems to be bounded to the 
west by the Erz Mountains, the Danube, the Alps, and the 
Rhone, all the specimens from east and south of that line be- 
longing to it, with a few exceptions mentioned below. All over 
France, west and north of the Rhone, the var. tceniata, described 
further on, alone occurs (with rare exceptions from the Doubs), 
whence it extends to ISTorthei'n Spain (Bilbao, _y?c/e Bedriaga) and 
Portugal (Oporto, Brit. Mus.). 



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MR. E. G. BOULENGER ON THE 









l^'k;^^;: 




All the specimens hitherto examined from Belgium, North- 
Western Germany, and the Rhine are referable to that variety, 
which also prevails in Wiirtemberg (cf. Leydig, 19). Exceptions 
to the above geographical division occur to my knowledge near, or 



VARIATIONS OF THE SrOTTED SALAMANUEU. 333 

not far from, the line of demarcation, as on Mt. Saleve in Savoy, 
according to M. Ghidini, but possibly also in Italy near Rome, 
where, according to Dumeril, the striped variety has been found 
(yellow on the back, with three black stripes, and a few scattered 
black spots on the limbs and belly) *. Further, a specimen 
referable to the typical form from Ballenstedt, Anhalt, received 
from Dr. Wolterstorff, one from the neighbourhood of Stuttgart, 
received from Prof. Lamport, and six exceptional specimens 
sketched by Mme. Phisalix : five from near Besangon, black 
above with irregular yellow blotches on the back, and one from 
Toulouse (Paris Museum), yellow above and beloAv with irregular 
black markings on the back, a very aberrant specimen. One of 
these specimens, from Beure, near Besangon, has been presented 
by Mme. Phisalix to the British Museum, and is listed under 
the head of forma typica, whilst six others from the same locality 
appear inider var. tceniata, thus showing that in the French 
Jura the latter variety is not so completely fixed as it appears 
to be in the North of France and Germany and Belgium. 
Five specimens are represented on text-fig. 101 : — 

a, fi-om Varese, Lombardy, is remai-kable for its small amoimt 

of yellow, and for its long and thin yellow vertebral stripe, 
extending from the nape to over halfway down the back. 
Sides and lower surface with only a few spots. 

b, from the Benider hills, Morocco, with a small number of 

large roundish spots on the back, has the left parotoid 
entirely black. 

c, from Hiitteldorf , near Yienna, has a remai-kably large number 

of spots. Flanks and sides of belly spotted. 

d, from the neighbourhood of Prague, is a type with very 

irregular markings. Flanks and sides of belly with only 
a few large spots. 

e, from Zebel, Bulgar Dagh, Asia Minor, has some of the spots 

ring-like, the yellow markings having a round black spot 
in the centre. Sides profusely spotted. Lower surface 
with many spots of small size. 

The specimen figured on Plate XV. is a male from Lugano, 
Ticino, sent alive by M. A. Ghidini. 

II. The Varieties of the Spaniibh Peninsula. 
(Vars. gallaica and 'molleri.) 

We have mentioned above that the var. tceniata occurs in 
Spain and Portugal. Specimens of that form, with the yellow 
colour predominating over the black, are stated to occur in the 
Peninsula by Schreiber (27, p. 78), and Bedriaga (3, p. 108) 

* I am indebted to Mme. Phisalix for a sketch of this specimen preserved in the 
Paris Museum. It is not unlikely, however, that the locality under which it 
has been registered is erroneous, as neither de Betta (4), Camerano (6), nor Count 
Peracca {in litt.) have come across the striped variety in any part of Italy. 



334 MR. E. G. BOULENGER ON THE 

records such a one from Bilbao. But in addition we find highly 
remarkable specimens ranging from tlie form named var. gallaica 
by Seoane (29), Avhich is hardly separable from the typical form, 
especially its North African representatives, to that named var. 
molleri by Bedriaga, which approaches very closely some speci- 
mens from nea,r Genoa, from Austria, described by Kammerer 
(c/. p. 342), and from Oran, Algeria, described by Doumergue *. 
My father has already proposed to unite the var. gallaica with the 
var. molleri, a view in which Bedriaga (3, p. 109) could not concur, 
on the ground that Seoane's diagnosis does not at all agree with his 
own. It is, nevertheless, a fact that an almost uninterrupted series 
can be traced between the two varieties, and I think it advisable, 
provisionally at least, to regard them as extremes of one and the 
same form, which is completely linked with the typical form. 

The following is a translation of Seoane's definition of the 
Spanish specimens (var. gallaica) : — " DiflJeis from the typical form, 
among other characters, in the intense black of the ground-colour 
and the small number of yellow spots, distributed over the 
body." 

The three specimens from Galicia [Seoane) in the Lataste 
Collection are remarkable for the very irregular, broken-up 
disposition of the spots on the back, a few of which are partly 
brownish and may have been edged with red or pink, in a manner 
similar to Bedriaga's var. molleri. The snout, supraocular and 
intei'orbital regions, partly yellow, partly reddish brown ; yellow 
on the throat somewhat predominating over the black ; sides 
irregularly spotted with yellow. The number of spots on the 
dorsal region of these specimens is 9, 13, and 26 respectively. A 
specimen from Cabanas, Galicia (Seoane), preserved in the Paris 
Museum and of which a sketch has kindly been made for me by 
Mme. Phisalix, approaches very closely the Portuguese var. 
molleri. The British Museum specimen from Vigo differs, how- 
ever, from all the above by being very scantily marked with 

* Essai sur la Fauiie Erp^tologique de I'Oranie (Oran, 1901), p. 372. 

" Corps presentant eu dessus plusieuvs taches jauiies et rouges sans symetvie 
doiit voici la distribution, liegions sus-oculaires jaunes en dessus et d'un rouge sang 
en avant et eu arriere. Arcades sourcilieres d'un noir rougeatre. Parotides jaunes 
en dessus et aussi en dessous posterieuremeut, entourees de noir en avant ; exte- 
rieureinent elles sont bordees depuis I'oBil jusque sur le cou, d'une loiigue et large 
taclie rouge. Seules les taches des regions sus-oculaires et celles des parotides pi'e- 
sentent quelque symetrie. Sur le cou se trouve une grande taclie transversale 
echancree en avant, a laquelle font suite, sur le dos, quatre taches irregulieres (de 7 
mill, sur 3 en moyenne). Ces taches alternent entre elles et touchent la double ligne 
dorsale de tubercules ; elles sont a peu pres a egale distance I'une de I'autre. Pres 
de I'aisselle, sur le bras, il y a une petite taclie jaune bordee de rouge ; une ou deux 
tres petites, jaunes et rouges, se voient sur Tavaiit bras, et une seule sur les mains et 
les pieds. Le fond noir des flancs est parsem^ de quelques points rouges. Membres 
post^rieures taches comme les ant(5rieurs. En arriere de la ligne des cuisses, en 
dessus, commence une tache jaune, longue et etroite, qui s'etend en arriere ; elle a 
10 mill, sur 2 a 3. Sur la queue ou voit cinq series de taches doubles, rondes, qui se 
rapprochent I'une de I'autre sur la ligne mediane superieure ; elles sont jaunes et 
visiblement bien bordees de rouge, surtout celle placees vers le bout de la queue. 
Mamelon du cloaque tach(5 de jaune de chaque cote. Dessous du corps d'un violet 
noiratre. Pourtour iuferieur de la bouche borde de taches rouges qui s'etendent sur 
la gorge." 



VARIATIONS OF THE SPOTTED SALAMANDEIJ. 335 

yellow, except on the gvilar region and on the sides of the belly. 
The back, snout, and interorbital region bear no distinct markings, 
but are speckled over with small yellowish dots. The parotoids 
and upper eyelids are almost entii'ely of a reddish-brown colour. 
This specimen must be I'egarrled as an individutd a.beriation of the 
form above described, an aberration tending to the total suppies- 
sion of the bright markings. 

Three specimens referable to the vai'. gallaica have been received 
from M, de la Escaleia, who obtained them in the Loroya Valley, 
near Madrid, at an altitude of 300 to 400 metres. Th.e spots are 
moderately large, few or moderately numerous (G to 12), those on 
the parotoids being either confluent with or distinct from those 
on the upper eyelids and the dorsal region. The spot on the 
eyelid in one of these specimens is entirely of a reddish brown, 
■chat on the parotoid partly reddish, partly yellow. In this and 
a.nother specimen the interorbital region is also reddish. The 
spot at the angle of the mouth in all three is brownish red in 
colour. Lower surface and sirles black, minutely speckled over 
with yellow; throat spotted with red. 

The true var. onolleri is represented in the British Museum 
Collection by 7 specimens from Poi-tugal (Coimbia and Cintra), 
including one of the types received from Dr. de Bedriaga, who 
has thus described its coloration : — 

" The colour and pattern of this vai'iety are rather variable. The 
ground-colour is usually a greyish blown, sometimes more of a 
dirty grey, sometimes more brownish black or even black, broken 
up above and below by pale j^ellow spots with an addition of grey 
or greyish-brown spots into which the yellow passes gradually. 
The yellow spots on the side of the body, on the limbs, on the tail, 
on the parotoids, on the tliroat, and on the eyelids are as if 
powdered over with red dust, or washed with red, or even blood- 
red. The throat may sometimes acquire a deep red colour ; the 
dorsal spots show hei-e and there a red dot. The yellow spots 
are A^ery variable both in number and size ; they may be either 
few, in which case they are large and roundish, or numerous and 
horseshoe- or ring-shaped, and forming six or eight more or less 
regular longitudinal series ; some of these spots break up or run 
together, thus forming wavy bands. These spots may be so 
iiumerous as to greatly reduce the ground-colour ; the yellow 
spots on the head are in that case the more conspicuous and a 
symmetrical or very ornamental pattern results." 

My father has drawn up the following notes on living specimens 
exhibited in our Zoological Gardens, a few years ago. 

" Some specimens were black, variegated with various tints of 
grey, brown, pale yellow, and crimson. The latter colour was 
particularly conspicuous on the upper eyelids, the parotoid glands, 
the base of the limbs, and on the throat, but it appeared also 
as small patches within the area of the more or less irregidar 
pale yellow spots with grey centres, which were disposed very 
irregularly on the body and tail. One of the specimens was pale 



336 MR. E. G. BOULENGER ON THE 

olive-grey above and on the sides, freckled with black and with 
pale greenish-yellow spots ; the black appeared as an irregular 
vertebral stripe, a dorso-lateral stripe, and bars on the flanks ; 
the upper eyelids, the parotoids, and the throat wei-e claret-red. 
The coloration of such a Salamander has a lichen-like aspect 
more suggestive of assimilation to the surroundings than of 
warning to enemies." Other specimens which he has seen since 
had but little or no red on them, but the yellow spots were 
greyish in the centre. 

In his description of var. molleri, Bedriaga states that it diifei's 
from the typical form in the snout being more pointed and 
projecting beyond the lower jaw, also that the tail is shorter and 
thicker. In five out of eight specimens (including Bedriaga's 
type) examined by me, I found the snout to be more projecting 
than is usual in the other forms, although the most pointed 
snout I have seen is in a specimen from near Meissen, Saxony. 
I also fovind that the tail in four out of the eight specimens 
was stouter and shorter than usual in the typical form and 
the var. tceniata, and this is also to be noticed in the figure 
on PL XY. The length of the tail in each of the specimens 
(measvired from the posterior end of the vent) was 55, 55, 57, 
57, 63, 67, 67, 71, the length of the body being taken as 100 (tip 
of snout to posterior end of vent). The length of the tail in the 
typical form varies from 60 to 78, with an average of about 67, 
whilst in the var. tceniata it may fall as low as 54 (Besan9on). 
It will be seen therefore, that there are many exceptions, and 
too much importance should not be attached to this character. 
I may add that the measurements of the Galician specimens, 
alluded to above under the name of var. gallaica, give 57, 66, 
70, and 70 as the length of the tail, and those of the specimens 
from near Madrid 56, 65, and 70. 

Thanks to the coui-tesy of the Hon. Walter Rothschild, I am 
able to give a figure (PI. XY.) of the var. molleri from a sketch 
made for him by Mr. J. Green, from a female specimen in his 
possession exhibited a few years ago in the Zoological Gardens. 

I have myself examined two fresh specimens of this variety 
from Lisbon, one alive, received from Sr. F. Mattozo Santos, 
Director of the Museum Bocage at Lisbon. In the live specimen 
the crimson-red colour was distributed over the parotoids, the 
upper eyelids, the throat, the spots at the angle of the mouth and 
on the sides, and the spot on the foi^earmand thigh. On the back 
and tail there was no red colour, but many of the yellow spots 
were partly bordered or as if washed over in the centre with a 
dirty grey. In the second specimen the red colour was restricted 
to the parotoids, the upper eyelids, and the spot at the angle of the 
mouth. The markings of the back, limbs, tail, sides, throat, and 
belly being mostly light grey in the centre and on the borders. 

On careful examination of the red and grey markings in these 
specimens, I have come to the conclusion that these are due, not 
to special pigments in addition to or in combination with the 



I 



VARIATIONS OF TIIK SPOTTED SALAMANDER. 337 

yellow, as 1ms been supposed, but: to the absence of pigment, the 
pigmentless Hesh, highly Unshed with blood, being exposed on 
certain patches and the grey colour resulting from the absence of 
yellow combined with a small quantity of black pigment. 

The lai-gest specimen of the var. molleri examined by me 
measures 109 mm. to the posterior extremity of the vent, the 
tail measuring 73. 

III. Var. t.^niata. 

This variety differs from the typical form in the arrangement 
of the dorsal spots, which are regularly disposed in two parallel 
series continuous with the patches on the parotoids, and not un- 
frequently form two uninterrupted stripes. Even when the tv.'0 
stripes are broken up into as many as 12 spots, these still retain 
their duplex disposition, not encroaching over the black vertebral 
line (Area A), or if they do so, as is rai'ely the case, only on the 
nape and the posterior end of the body, where they may be con- 
nected in H-like fashion by a cross-bar. Although in this variety 
the black often predominates over the yellow, it is not uiicommon 
to find specimens in which the reverse takes place. In cases 
where the yellow has so far invaded the upper surface as to actually 
constitute the ground-colour, the black vertebral stripe may be 
reduced to a mere series of spots, and Werner (31, p. 155, pi. vii, 
fig. 23) even mentions and figures one in which the black is com- 
pletely absent from the back. The upper eyelid and the parotoid 
gland are entirely yellow (the yellow patch on the latter raxely 
broken up), the two spots nearly always running together and 
often also confluent with the markings on the back, which is rarely 
the case with the typical form. The sides are usually devoid of 
markings, and if present they only occur in small numbers. 
Yellow markings are usually present on the snout, which is but ex- 
ceptionally the case with the typical form. The spot situated at the 
angle of the mouth is absent in about 35 per cent, of the specimens. 
Gular region and lower surface of body with a varying amount of 
yellow, sometimes entirely yellow with a, black spot or bar on the 
gular fold ; the markings have often a tendency to dispose them- 
selves in longitudinal series, thus the belly may be black with 
a yellow lateral stripe or yellow with one or two black stripes in 
the middle. Limbs with the black usually predominating ; a 
characteristic yellow blotch near the base of the arm and thigh, 
and a second on the forearm and leg being constant, and usually 
larger than in the typical form. Hand and foot mostly black and 
yellow, the yellow patches being nearly always confluent with those 
on the forearm and leg. Nearly entirely yellow specimens have 
likewise yellow limbs M-ith merely 2 or 3 black spots or cross-bars. 
As on the limbs, the yellow may predominate over the black on the 
tail and frequently extend to the lower surface, which is rai'ely 
the case in the typical form. The two yellow dorsal bands often 
fuse on the upper surface of the tail. 

The colour varies from sulphur- or lemon-yellow to a deep orange. 



338 MR. E. G. BOULENGER ON THE 

Some speemiens with the markings almost vermilion-red, instead 
of yellow or orange, obtained by Fr. v. Schweizerbath near 
Stuttgart, are regarded by her as a distinct variety and named 
var. coccinea (28), but this is clearly to be regarded as a merely 
individual peculiarity, not deserving of a varietal name, and the 
figure given by her corresponds, but for the colours of the markings, 
with the var. tceniata, the prevalent form round Stuttgart. Con- 
sidering that the bright markings may vary, in the same locality, 
from chrome-yellow to a deep orange, the so-called var. coccinea 
represents merely an intensification of a tendency existing in 
German specimens. Fr. v. Schweizei-bath has been informed by 
Prof. E. Haeckel that such a Salamander was found by him many 
years ago in the Saal Valley, near Ziegenriick, and it is not un- 
likely that vermilion-spotted specimens will be discovered in other 
parts of the habitat of the var. tceniata. My father was informed 
by an intelligent peasant woman in Belgium that on the occasion 
of her witnessing, in a wood, just before a thunderstorm, a sudden 
apparition of Salamanders in huge numbers, some among them 
were distinguished by being marked with red instead of yellow. 
In D'Orbigny's ' Dictionnaire d'Histoire Naturelle ' (7, p. 307) 
allusion is made to a specimen found near Bordeaux, which must 
have been similar to the one described by Fr. v. Schweizerbath. 
In Belgium, the mai-kings are of a more or less bright yellow but 
not orange, and in most cases they form interrupted stripes. In 
Brittany the markings vary from sulphur- to chrome-yellow. Out 
of 50 specimens obtained together, within a space of one hundred 
square yards, last summer at RoscofF, about half had the stripes 
uninterrupted but varying much in width. M. Ghidini, of the 
Geneva Museum, having had occasion to examine 500 specimens 
received alive from Stadtoldendorf in Brunswick, found that 
about 400 had the two parallel stripes uninterrupted, or nearly 
so, 50 had them much bioken up, whilst the remaining 50 were 
nearly entirely yellow, with the black reduced to spots or narrow 
stripes. The specimens from the Harz and neighbouring hills, of 
which I have seen many, vary in the colour of the markings from 
chrome-yellow to a rather deep orange. 

A male specimen from the Harz, in Avhich the yellow colour 
prevails, is represented on Plate XY. 

Figures of the var. tceniata are given by "VVurf bain (33), Gesner 
(13, ii. p. 80), Perrault (21, pk 16. p. 77), Duvernoy (11, pi. xl. 
fig. 1), Mme. Phisalix (22, pi. i.), and Diirigen(10, p. 577). The 
descriptions of Leydig (19), Lataste (16), and Martin and Rollinat 
(20) are also applicable to it, as well as the var. B of Dumeril and 
Bibron (9) and the vars./to k of Schreiber (27). 

Five specimens are represented in text-fig. 102, to give so